一株高致病性鸡传染性贫血病毒的生物学及基因组特性
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摘要
鸡传染性贫血病毒(Chicken anemia virus, CAV)在世界主要养禽国家广泛分布,是危害养禽业的主要病原体之一。CAV感染可以破坏造血系统,并可使淋巴器官萎缩,导致鸡群免疫抑制,这又经常使感染鸡群并发或继发其它病原体的多重感染,导致巨大经济损失。因此对CAV的系统研究甚为重要,本研究是从CAV的检测方法、流行情况以及流行毒株的基因变异和致病性进行系统的研究。
根据GenBank中已经发表的CAV VP1基因的序列,设计合成一对引物,利用PCR技术扩增CAV VP1基因的的核酸片段,利用非放射性标记物地高辛(DIG),通过PCR方法制备了CAV核酸探针。通过斑点杂交检测方法对此探针特异性及灵敏度进行验证,特异性检测结果表明,该探针能与标准模式株CAV核酸发生特异性杂交,而与对照的SPF鸡、禽网状内皮组织增生症病毒(REV)、马立克氏病病毒(MDV)、呼肠孤病毒(REOV)核酸杂交反应为阴性,具有较强的特异性。敏感性检测结果表明,该探针对CAV的最低检出量为1 pg核酸,具有较高的灵敏度,显示所制备的核酸探针用于临床病料组织DNA中CAV感染的检测是可行的。
本实验收集山东泰安、临沂、莱芜、烟台等8市3个品系、不同日龄发病鸡群的384只病、死鸡的脾脏样品,其中麻鸡105份、肉鸡185份、蛋鸡94份,通过斑点杂交方法检测样品中的CAV核酸,检测结果为,CAV总检出率为27.9%(107/384)。3个品系的鸡的检测结果:麻鸡样品的检出率为37.1%(39/105);肉鸡样品的检出率为31.3%(58/185);蛋鸡样品的检出率11.7%(11/94)。结果显示,麻鸡和肉鸡中CAV感染的检出率显著高于蛋鸡。3个不同日龄段的检测结果:1-20日龄的样品检出率为6.4%(7/110);20-60日龄的样品检出率为45.6%(93/204);60日龄以上的样品检出率为10.0%(7/70)。结果显示,20-60日龄的检出率显著高于1-20日龄和60日龄以上的,而20-60日龄段是疾病的高发阶段。
其中,从自山东某商品代肉鸡场20日龄病鸡群表现生长迟缓,疑似鸡传染性贫血病病毒(CAV)感染的脾脏、胸腺、骨髓样品提取DNA,分别经斑点杂交检测为CAV阳性后,其余病料研磨上清经处理后接种7日龄SPF鸡胚和1日龄SPF鸡,提取接种后的鸡胚和SPF鸡的脾脏组织DNA,经斑点杂交验证为阳性,从而成功分离到了一株鸡传染性贫血病毒野毒株,命名为CAV-SDLY08株。用PCR方法分段扩增出CAV-SDLY08基因组的三条部分重叠片段,分别克隆于T载体并进行测序,拼接后得到其全基因组序列。结果表明,SDLY08株基因组全长2298nt,含有三个互相重叠的开放阅读框和一个调控区。将SDLY08全基因与已发表的8个CAV分离株基因组比较,同源性为96.6%-99.0%。CAV的三个编码基因VP1、VP2和VP3所编码的氨基酸均有一定程度变异,以VP1基因编码的氨基酸变异性最大,同源性为96.9%-99.0%
将CAV-SDLY08株接种感染1、7、21日龄SPF鸡,研究其致病性。通过口服和肌肉注射两种途径分别感染1、7、21日龄SPF鸡,12天后对每组SPF鸡体重、免疫器官指数和血液指标进行测定。
体重增重方面:感染12天后,1、7日龄感染组的体重显著低于对照组(P<0.05),而两不同途径感染组间的差异不显著(P>0.05)。21日龄感染组体增重也低于对照组,但差异不显著(P>0.05),两种感染方式的差异也不显著。
免疫器官指数方面:感染12天后,不同日龄的感染均能引起脾脏的肿大,1日龄感染的口服组和注射组与对照组比差异显著(P<0.05)。7日龄和21日龄的与对照组比,差异不显著(P>0.05),且感染途径影响不大。3个日龄感染都能引起胸腺的显著萎缩,且不同的感染日龄和感染方式的实验组与对照组相比差异都极显著(P<0.01)。
血液指标方面:感染12天后,可引起白细胞数量的减少,1日龄感染的口服组和注射组与对照比差异极显著(P<0.01)。7日龄和21日龄,注射组显著低于口服组和对照组(P<0.05),口服组和对照组差异不显著(P>0.05)。与对照组相比,CAV感染都能引起红细胞数量的显著减少(P<0.01),口服组和注射组与对照组相比差异极显著(P<0.01),且两种感染方式之间差异显著(P<0.05)。与对照组相比,都能引起红细胞压积的显著减少(P<0.01),1日龄和7日龄感染的口服组和注射组差异极显著(P<0.01),21日龄差异显著(P<0.05)。
综合结果显示, CAV-SDLY08株感染三个日龄SPF鸡12天后,均可导致增重减缓、胸腺显著萎缩、脾脏肿大、白细胞数、红细胞数与红细胞压积的显著减少。这表明该CAV-SDLY08分离株对1日龄和7日龄鸡的致病性要强于21日龄,肌肉注射比口服感染组的致病性更强。而且结果同时表明,该毒株对21日龄的SPF鸡仍有明显的致病作用。
Chicken anemia virus (CAV) is widely distributed in the world and is one the most important pathogens in chiken industry. CAV causes transient severe aplastic anemia due to destruction of erythroblastoid cells and generalized lymphoid atrophy with a concomitant immunosuppression.CAV causes considerable economic losses in poultry industry as a result of a co-infection and a secondary affection after its infection. So it is important to reaserch CAV systematically.In this study,we systematically Research CAV in aspects of Detection methods,epidemiology and genetic variation and pathogenicity of a pandemic strain.
According to the genomic sequences of VP1 gene of Chicken anemia virus (CAV) published in Genbank,one pairs of primers were designed for amplifying the fragment in PCR experiments. The PCR product was labeled with digoxigenin by PCR as DNA probe for detection of CAV. The dot blot hybridization assay result of specificity showed CAV was positive, but other nucleotide extracted from REV,MDV,REOV were negative. The sensitivity result showed that as few as 1 pg DNA amount of CAV could be detected by DIG-labeled probe.So the DIG-labeled probe could be used to detect the CAV.
The spleen samples of 384 chickens with clinical performance from 3 strains chicken flocks of different ages were collected in eight cities of Shan dong province, including 105 portion partridge chickens,185 portion broilers,94 portion egg chickens. By dot blot hybridization assay , the detection result of CAV in 384 portion spleen samples showed that total detection rate was 27.9%(107/384). Test results of 3 strains chickens showed that detection rate of partridge chickens was 37.1%(39/105); detection rate of broilers was 31.3%(58/185);detection rate of egg chickens was 11.7%(11/94). Tthe experiments indicated that detection rate of broilers and partridge chickens was significantly higher than egg chikens.Test results of 3 ages chickens showed that detection rate of 1-20 old-day ages was 6.4%(7/110); detection rate of 20-60 old-day ages was 45.6%(93/204);detection rate of after 60 old-day ages was 10.0%(7/70). The experiments indicated that detection rate of 20-60 old-day ages was significantly higher than 1-20 old-day ages and after 60 old-day ages, 20-60 old-day ages was the peak disease prases.
Extract DNA from spleen、thymus and bone marrow of disease chicken suspected infection with CAV from a 20 day-old broiler flock with growth runting syndromes, dot blot hybridization showed CAV positive, A field strain SDLY08 of chicken infectious anemia virus (CAV) was isolated by 1 day-old Specific-pathogen-free(SPF) chicken and 7 day-old Specific-pathogen-free (SPF) chicken embryo inoculation. The viral genomic DNA was amplified by PCR in 3 overlapped fragments and PCR products were cloned into T-vecor plasmid for sequencing. The sequencing results indicated that the total genome of SDLY08 strain CAV was 2298nt, it contained 3 overlapped ORF and 1 non-coding regulation fragment. Its whole genome had 96.6%-99.0% of homogeneity to other 8 published CAV reference strains. There were some mutations in 3 amino acid sequence which were encoded by 3 genes VP1, VP2 and VP3. VP1 was less conservative than VP2 and VP3. the homogeneity was 96.6%-99.0%.
In this study, Specific-pathogen-free(SPF) chickens were inoculated of a chicken infectious anemia virus field isolate CAV-SDLY08 with 104.5 mean embryo infective dose(EID50) at 1、7、21 day of age to elucidate the pathogenicity of CAV- SDLY08.SPF chickens at different ages of 1、7、21 day were challenged with the virus by oral administration and intramuscular injection. Ratios of thymuses, spleens and the Bursa to body weight, several blood parameters were examined and compared to controls 12 days after infectons with CAV.
The reaserch of weight show that the weight of 1-day-old and 7-day-old chickens was significantly lower than control group (P<0.05) 12 days after infectons with CAV,the differences in two infective methods were not significant statistically(P>0.05). The infective group of 21-day-old chickens was also lower than control group,but differences were not significant statistically(P>0.05) ,the differences in two infective methods were not significant statistically.
The reaserch of immune organ show that there was splenomegaly in different ages, the differences were significant statistically in the oral administration group and the injection group of 1-day-old chickens compared to control group(P<0.05). The differences were not significant statistically in 7-day-old chickens group and 21-day-old chickens group compared to to control group(P>0.05).There was no differences in two infective methods.The infection of 3 ages could all cause severe atrophy of thymus, the differences were very significant statistically in different infective ages and different infective methods compared to control group(P<0.01).
The reaserch of blood index show that, white blood cells decrease after infection of 12 day, the differences were very significant statistically in the oral administration group and the injection group of 1-day-old chickens compared to control group(P<0.01). With infection of 7-day-old chickens group and 21-day-old chickens group, the injection group was significantly lower than oral administration group and control group (P<0.05),there was no significant differences in oral administration group and control group (P>0.05).Infection of CAV strain SDLY08 at all 3 ages could cause significantly decrease of red blood cells, the differences were very significant statistically in the oral administration group and the injection group compared to control group(P<0.01) and the differences of two methods infection were significant statistically(P<0.05). The haematocrit decrease significantly compared to control group(P<0.01), the differences were very significant statistically in the oral administration group and the injection group of 1-day-old chickens and 7-day-old chickens compared to control group(P<0.01), significant in 21-day-old chickens (P<0.05).
Results indicated that challenges of CAV strain SDLY08 at all 3 ages could cause growth retardation, severe atrophy of thymus and spleen megaly, significant decrease of red blood cells、white blood cells and the haematocrit. It was also demonstrated that 1-day-old and 7-day-old SPF chickens were more sentitive to SDLY08 than 21-day-old chickens, chickens were more sensitive to challenges by intramuscular injection than oral administration. Results also showed that this CAV strain can also cause significant pathogenicity in 21-day-old chickens
根据GenBank中已经发表的CAV VP1基因的序列,设计合成一对引物,利用PCR技术扩增CAV VP1基因的的核酸片段,利用非放射性标记物地高辛(DIG),通过PCR方法制备了CAV核酸探针。通过斑点杂交检测方法对此探针特异性及灵敏度进行验证,特异性检测结果表明,该探针能与标准模式株CAV核酸发生特异性杂交,而与对照的SPF鸡、禽网状内皮组织增生症病毒(REV)、马立克氏病病毒(MDV)、呼肠孤病毒(REOV)核酸杂交反应为阴性,具有较强的特异性。敏感性检测结果表明,该探针对CAV的最低检出量为1 pg核酸,具有较高的灵敏度,显示所制备的核酸探针用于临床病料组织DNA中CAV感染的检测是可行的。
本实验收集山东泰安、临沂、莱芜、烟台等8市3个品系、不同日龄发病鸡群的384只病、死鸡的脾脏样品,其中麻鸡105份、肉鸡185份、蛋鸡94份,通过斑点杂交方法检测样品中的CAV核酸,检测结果为,CAV总检出率为27.9%(107/384)。3个品系的鸡的检测结果:麻鸡样品的检出率为37.1%(39/105);肉鸡样品的检出率为31.3%(58/185);蛋鸡样品的检出率11.7%(11/94)。结果显示,麻鸡和肉鸡中CAV感染的检出率显著高于蛋鸡。3个不同日龄段的检测结果:1-20日龄的样品检出率为6.4%(7/110);20-60日龄的样品检出率为45.6%(93/204);60日龄以上的样品检出率为10.0%(7/70)。结果显示,20-60日龄的检出率显著高于1-20日龄和60日龄以上的,而20-60日龄段是疾病的高发阶段。
其中,从自山东某商品代肉鸡场20日龄病鸡群表现生长迟缓,疑似鸡传染性贫血病病毒(CAV)感染的脾脏、胸腺、骨髓样品提取DNA,分别经斑点杂交检测为CAV阳性后,其余病料研磨上清经处理后接种7日龄SPF鸡胚和1日龄SPF鸡,提取接种后的鸡胚和SPF鸡的脾脏组织DNA,经斑点杂交验证为阳性,从而成功分离到了一株鸡传染性贫血病毒野毒株,命名为CAV-SDLY08株。用PCR方法分段扩增出CAV-SDLY08基因组的三条部分重叠片段,分别克隆于T载体并进行测序,拼接后得到其全基因组序列。结果表明,SDLY08株基因组全长2298nt,含有三个互相重叠的开放阅读框和一个调控区。将SDLY08全基因与已发表的8个CAV分离株基因组比较,同源性为96.6%-99.0%。CAV的三个编码基因VP1、VP2和VP3所编码的氨基酸均有一定程度变异,以VP1基因编码的氨基酸变异性最大,同源性为96.9%-99.0%
将CAV-SDLY08株接种感染1、7、21日龄SPF鸡,研究其致病性。通过口服和肌肉注射两种途径分别感染1、7、21日龄SPF鸡,12天后对每组SPF鸡体重、免疫器官指数和血液指标进行测定。
体重增重方面:感染12天后,1、7日龄感染组的体重显著低于对照组(P<0.05),而两不同途径感染组间的差异不显著(P>0.05)。21日龄感染组体增重也低于对照组,但差异不显著(P>0.05),两种感染方式的差异也不显著。
免疫器官指数方面:感染12天后,不同日龄的感染均能引起脾脏的肿大,1日龄感染的口服组和注射组与对照组比差异显著(P<0.05)。7日龄和21日龄的与对照组比,差异不显著(P>0.05),且感染途径影响不大。3个日龄感染都能引起胸腺的显著萎缩,且不同的感染日龄和感染方式的实验组与对照组相比差异都极显著(P<0.01)。
血液指标方面:感染12天后,可引起白细胞数量的减少,1日龄感染的口服组和注射组与对照比差异极显著(P<0.01)。7日龄和21日龄,注射组显著低于口服组和对照组(P<0.05),口服组和对照组差异不显著(P>0.05)。与对照组相比,CAV感染都能引起红细胞数量的显著减少(P<0.01),口服组和注射组与对照组相比差异极显著(P<0.01),且两种感染方式之间差异显著(P<0.05)。与对照组相比,都能引起红细胞压积的显著减少(P<0.01),1日龄和7日龄感染的口服组和注射组差异极显著(P<0.01),21日龄差异显著(P<0.05)。
综合结果显示, CAV-SDLY08株感染三个日龄SPF鸡12天后,均可导致增重减缓、胸腺显著萎缩、脾脏肿大、白细胞数、红细胞数与红细胞压积的显著减少。这表明该CAV-SDLY08分离株对1日龄和7日龄鸡的致病性要强于21日龄,肌肉注射比口服感染组的致病性更强。而且结果同时表明,该毒株对21日龄的SPF鸡仍有明显的致病作用。
Chicken anemia virus (CAV) is widely distributed in the world and is one the most important pathogens in chiken industry. CAV causes transient severe aplastic anemia due to destruction of erythroblastoid cells and generalized lymphoid atrophy with a concomitant immunosuppression.CAV causes considerable economic losses in poultry industry as a result of a co-infection and a secondary affection after its infection. So it is important to reaserch CAV systematically.In this study,we systematically Research CAV in aspects of Detection methods,epidemiology and genetic variation and pathogenicity of a pandemic strain.
According to the genomic sequences of VP1 gene of Chicken anemia virus (CAV) published in Genbank,one pairs of primers were designed for amplifying the fragment in PCR experiments. The PCR product was labeled with digoxigenin by PCR as DNA probe for detection of CAV. The dot blot hybridization assay result of specificity showed CAV was positive, but other nucleotide extracted from REV,MDV,REOV were negative. The sensitivity result showed that as few as 1 pg DNA amount of CAV could be detected by DIG-labeled probe.So the DIG-labeled probe could be used to detect the CAV.
The spleen samples of 384 chickens with clinical performance from 3 strains chicken flocks of different ages were collected in eight cities of Shan dong province, including 105 portion partridge chickens,185 portion broilers,94 portion egg chickens. By dot blot hybridization assay , the detection result of CAV in 384 portion spleen samples showed that total detection rate was 27.9%(107/384). Test results of 3 strains chickens showed that detection rate of partridge chickens was 37.1%(39/105); detection rate of broilers was 31.3%(58/185);detection rate of egg chickens was 11.7%(11/94). Tthe experiments indicated that detection rate of broilers and partridge chickens was significantly higher than egg chikens.Test results of 3 ages chickens showed that detection rate of 1-20 old-day ages was 6.4%(7/110); detection rate of 20-60 old-day ages was 45.6%(93/204);detection rate of after 60 old-day ages was 10.0%(7/70). The experiments indicated that detection rate of 20-60 old-day ages was significantly higher than 1-20 old-day ages and after 60 old-day ages, 20-60 old-day ages was the peak disease prases.
Extract DNA from spleen、thymus and bone marrow of disease chicken suspected infection with CAV from a 20 day-old broiler flock with growth runting syndromes, dot blot hybridization showed CAV positive, A field strain SDLY08 of chicken infectious anemia virus (CAV) was isolated by 1 day-old Specific-pathogen-free(SPF) chicken and 7 day-old Specific-pathogen-free (SPF) chicken embryo inoculation. The viral genomic DNA was amplified by PCR in 3 overlapped fragments and PCR products were cloned into T-vecor plasmid for sequencing. The sequencing results indicated that the total genome of SDLY08 strain CAV was 2298nt, it contained 3 overlapped ORF and 1 non-coding regulation fragment. Its whole genome had 96.6%-99.0% of homogeneity to other 8 published CAV reference strains. There were some mutations in 3 amino acid sequence which were encoded by 3 genes VP1, VP2 and VP3. VP1 was less conservative than VP2 and VP3. the homogeneity was 96.6%-99.0%.
In this study, Specific-pathogen-free(SPF) chickens were inoculated of a chicken infectious anemia virus field isolate CAV-SDLY08 with 104.5 mean embryo infective dose(EID50) at 1、7、21 day of age to elucidate the pathogenicity of CAV- SDLY08.SPF chickens at different ages of 1、7、21 day were challenged with the virus by oral administration and intramuscular injection. Ratios of thymuses, spleens and the Bursa to body weight, several blood parameters were examined and compared to controls 12 days after infectons with CAV.
The reaserch of weight show that the weight of 1-day-old and 7-day-old chickens was significantly lower than control group (P<0.05) 12 days after infectons with CAV,the differences in two infective methods were not significant statistically(P>0.05). The infective group of 21-day-old chickens was also lower than control group,but differences were not significant statistically(P>0.05) ,the differences in two infective methods were not significant statistically.
The reaserch of immune organ show that there was splenomegaly in different ages, the differences were significant statistically in the oral administration group and the injection group of 1-day-old chickens compared to control group(P<0.05). The differences were not significant statistically in 7-day-old chickens group and 21-day-old chickens group compared to to control group(P>0.05).There was no differences in two infective methods.The infection of 3 ages could all cause severe atrophy of thymus, the differences were very significant statistically in different infective ages and different infective methods compared to control group(P<0.01).
The reaserch of blood index show that, white blood cells decrease after infection of 12 day, the differences were very significant statistically in the oral administration group and the injection group of 1-day-old chickens compared to control group(P<0.01). With infection of 7-day-old chickens group and 21-day-old chickens group, the injection group was significantly lower than oral administration group and control group (P<0.05),there was no significant differences in oral administration group and control group (P>0.05).Infection of CAV strain SDLY08 at all 3 ages could cause significantly decrease of red blood cells, the differences were very significant statistically in the oral administration group and the injection group compared to control group(P<0.01) and the differences of two methods infection were significant statistically(P<0.05). The haematocrit decrease significantly compared to control group(P<0.01), the differences were very significant statistically in the oral administration group and the injection group of 1-day-old chickens and 7-day-old chickens compared to control group(P<0.01), significant in 21-day-old chickens (P<0.05).
Results indicated that challenges of CAV strain SDLY08 at all 3 ages could cause growth retardation, severe atrophy of thymus and spleen megaly, significant decrease of red blood cells、white blood cells and the haematocrit. It was also demonstrated that 1-day-old and 7-day-old SPF chickens were more sentitive to SDLY08 than 21-day-old chickens, chickens were more sensitive to challenges by intramuscular injection than oral administration. Results also showed that this CAV strain can also cause significant pathogenicity in 21-day-old chickens
引文
[1]崔现兰,辛贵香,吴冬来.鸡传染性贫血抽毒的鉴定.中国畜禽传染病,1992,(6):3-5.
[2]殷震,刘景华.动物病毒学[M].2版.北京:科学出版社.1997.
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[3]崔治中,孟珊珊,姜世金,等.我国白羽肉用型鸡群中CAV、REV和REOV感染状况的血清学调查[J].畜牧兽医学报,2006,37(2):152-157.
[4]梁智选,杨汉春.我国部分地区鸡贫血病病毒全基因组核苷酸序列的比较与分析.中国预防兽医学报,2008,30(2):127-131.
[5]李延鹏,崔治中.一株鸡传染性贫血病毒野毒株致病性及其全基因组序列比较[J].微生物学报,2007,47(5):894-898.
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[7]崔治中.免疫抑制性病毒多重感染在鸡群疫病发生和流行中的作用[J].畜牧兽医学报,2003,34(5):417-421.
[8]崔治中.禽病诊治彩色图谱[M].北京:中国农业出版社.2004.
[9]崔治中.单股环DNA病毒基因组的复制及其转录调控因子结合序列.生命的化学,1995,15(3):23-25.
[10]何成强,丁乃峥,李景鹏等.鸡贫血病毒哈尔滨分离株全基因克隆和序列分析.微生物学报,2002,42(4):436-441.
[11]姜世金,孟珊珊,崔治中,等.我国自然发病鸡中MDV、REV和CAV共感染的检测[J].中国病毒学,2005,20(2):164-167.
[12]金文杰,崔治中,等.传染性法氏囊病病料中MDV、CAV、REV的共感染检测[J].中国兽医学报,2001,21:6-9.
[13]刘岳龙,崔治中,段玉友.PCR和斑点杂交检测鸡传染性贫血病毒.中国兽医学报,1996,16(1): 38-41.
[14]刘忠贵,郑世民,杨丽萍,等.感染鸡贫血病毒雏鸡接种新城疫疫苗后免疫保护效应及其机理研究[J].中国农业科学, 2001,34(1):1-4.
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[16]韦莉.鸡传染性贫血病毒分子生物学进展.北京农业科学, 1999,17(4):31-33.
[17]熊永忠,王秀荣,王笑梅.鸡传染性贫血病流行特点与诊断方法.黑龙江畜牧兽医,2002,(9):27-28.
[18]徐彦波,金华,甘永华等.鸡贫血病毒感染雏鸡细胞周期的研究.中国预防兽医学报,1999,21(2):122-123.
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[21]李延鹏,崔治中,等.CAV与REV共感染SPF鸡对疫苗免疫反应的抑制作用[J].中国兽医学报,2007,11:1243-1246.
[22] McNulty M S. Chicken anemia agent: a review. Avian Pathology,1991, 20:187-203.
[23] McNulty, M. S., T. J. Connor, F. McNeilly et al. Chicken anemia agent in the United States: Isolation of the virus and detection of antibody in broiler breeder flocks. Avian Dis,1989,33:691-94.
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[25] Yuasa N T,Taniguchi and Yoshida I. Isolation and some properties of an agent inducing anemia in chicks[J]. Avian Dis,1979,23:366-385.
[26] McNulty,M.S.,W.L.Curran,D.Todd,et al.Chicken anemia agent:An elec- tron microscopic study. Avian Dis.1990,34:736-743.
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[29] Zhou W,Shen B,Yang B,et al.Isolation and identification of chickeninfectious anemia virus in China[J].Avi.Dis.,1997,41(2):361-364.
[30] Yuasa N, T Taniguchi, T Noguchi, et al.Effect of infectious bursal disease virus infection on incidence of anemia by chicken anemia agent. Avian Diseases, 1980, 24:202-209.
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