氢溴酸常山酮抗柔嫩艾美耳球虫作用研究
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摘要
球虫病是由顶复门细胞内寄生性原虫——艾美耳球虫引起的一种鸡肠道寄生虫病。球虫病对养鸡业危害非常严重,因死亡率较高,病愈鸡饲料报酬下降、生长缓慢、产蛋率下降以及防治用的药物和疫苗的使用等原因,每年给世界养禽业造成约三十亿美元的经济损失。目前,使用化学药物防治是控制鸡球虫病的主要策略。化学药物能够预防该病的爆发,但亚临床感染相当普遍,而且球虫非常容易产生耐药性,新药研制费时费力,远远跟不上耐药性产生的速度。目前国内外许多学者在筛选各种抗球虫药物,但一直没有令人满意的结果。因此,从现有的药物入手,在研究其作用机理的基础上再进行新药的研发,或许是一条好的途径。氢溴酸常山酮自上个世纪三十年代以来就一直在使用,但对于其作用机理的研究却相对较少,对其作用的靶点不甚明了。因此,我们对氢溴酸常山酮抗柔嫩艾美耳球虫的作用开展了研究。
一、华东部分地区肉鸡球虫病的流行病学调查
为了了解华东地区肉鸡球虫病的流行情况,更好地为鸡球虫病的防治提供科学的依据,本项目组成员2011年3、4月份采集了来自华东地区的浙江省、江苏省、上海市、安徽省、山东省、江西省的养鸡场的429份鸡粪样品,进行了球虫卵囊的分离与鉴定。结果表明,有116份阳性样品,多数样品为两三种球虫的混合感染,六省市球虫病平均感染率为27.1%,江西、上海、山东、江苏四省市的球虫病感染率为30.0%-43.3%,高于平均水平。经单卵囊繁殖和多重PCR鉴定表明,上述六省市主要流行的球虫种类有四种,即巨型艾美耳球虫、柔嫩艾美耳球虫、毒害艾美耳球虫和堆型艾美耳球虫。
二、氢溴酸常山酮的抗球虫效果研究
氢溴酸常山酮是一类化学合成的高效广谱抗球虫药物,为了验证该产品对鸡球虫的作用效果,进行了本试验。本试验对14日龄鸡进行人工感染柔嫩艾美耳球虫,并用1.5mg/kg、3mg/kg和6mg/kg三种浓度的氢溴酸常山酮进行治疗。结果表明,与阳性对照组相比,1.5mg/kg、3mg/kg可以使球虫感染鸡的体重显著提高。随着药物浓度增加,没有观察到血便排出,没有观察到盲肠病变,但卵囊排出量和体重相对增重率都呈下降趋势。3mg/kg氢溴酸常山酮抗球虫指数高达195.45,表明该氢溴酸常山酮具有高效抗球虫效果。
三、氢溴酸常山酮的安全性评价
本实验以高于使用剂量1-3倍连续3周给药,通过鸡的增重情况、血液学与血液化学指标及组织病理学检查,评价了氢溴酸常山酮对鸡的安全性。实验结果表明,各实验组与对照组的血液化学指标和血液学指标无显著差异,尸体解剖未见各组织器官有明显的异常变化,组织学检查也未见有病理变化,说明氢溴酸常山酮对鸡只比较安全。
四、氢溴酸常山酮的体内作用机理研究
氢溴酸常山酮作为抗球虫药已有多年的使用历史,但是人们对其作用的机理却了解不多。为了研究其作用机理,本研究中分别对人工感染柔嫩艾美耳球虫后0、1、2、3、4、5、6天的鸡用3mg/kg饲料的氢溴酸常山酮进行治疗(相应的分别为第0、1、2、3、4、5、6组),观察不同给药时间的抗球虫效果。第7组的鸡只感染柔嫩艾美耳球虫而不治疗,作为感染对照组,第8组的鸡不感染不治疗,作为阴性对照组。感染后观察记录各组鸡的死亡率、体重增重率、卵囊下降率和盲肠病变记分,并且采集盲肠组织制备组织切片,综合评价氢溴酸常山酮的作用效果,初步探索其作用机理。实验结果表明,感染的当天或者感染后2天之内用氢溴酸常山酮治疗是完全有效的,如果感染2天后再用氢溴酸常山酮治疗只能起到部分保护效果或无保护效果。盲肠的组织切片观察表明,氢溴酸常山酮可以抑制柔嫩艾美耳球虫子孢子的入侵,氢溴酸常山酮治疗组的第二代裂殖体表现出空泡化和降解。因此,氢溴酸常山酮主要作用于球虫生活史的早期,阻止其入侵宿主细胞,然后通过抑制裂殖体分裂为裂殖子,使裂殖体空泡化达到破坏其发育的作用效果,导致裂殖体和裂殖子发育被抑制,发育速度减慢,从而被机体的免疫系统清除。
五、氢溴酸常山酮的体外作用机理研究
为了探索氢溴酸常山酮的体外作用机理,本实验对获得的新鲜的未孢子化柔嫩艾美耳球虫卵囊通过孢子化不同时间后给予氢溴酸常山酮孵育处理,再继续进行孢子化,观察了氢溴酸常山酮在体外条件下对孢子化的影响、卵囊的外观形态和超微结构的变化及部分有代表性关键基因的表达情况,并研究了子孢子体外处理后对细胞感染能力的影响。结果表明,氢溴酸常山酮可以抑制孢子生殖,使卵囊孢子化率降低,并且药物处理后的卵囊脱孢子囊能力下降,卵囊表面形态和细胞质及其超微结构等都发生明显异常,而且子孢子与氢溴酸常山酮体外条件下共孵育后对细胞的感染能力下降。根据结果推测,氢溴酸常山酮可能对卵囊壁的结构具有破坏作用。对多个基因的表达情况分析表明,氢溴酸常山酮作用后,丙醇酸激酶基因的表达下调,而1-磷酸甘露醇脱氢酶、微线体蛋白1和热激蛋白90基因表达上调。
综上所述,本研究对华东地区部分省市的鸡球虫病进行了流行病学调查,并用形态学方法和多重PCR方法对分离到的卵囊进行了虫种的鉴定；对氢溴酸常山酮进行了抗球虫效果实验及对鸡的安全性实验,并通过氢溴酸常山酮在球虫感染鸡体后不同时间给药探讨了抗柔嫩艾美耳球虫的作用机理,还在体外条件下,研究了氢溴酸常山酮对孢子生殖的影响,初步阐述了氢溴酸常山酮潜在的作用靶点及作用机理,但这些初步的结论还需要进一步的深入研究进行验证,这可能为球虫病的防控及药物开发提供借鉴意义。
Coccidiosis is caused by the intracellular protozoal Eimeria genus, which parasite in chicken's intestinal tract. Coccidiosis causes huge impaire to the poultry industry about$30billion annually because of the high mortality, reduced feed convertion rate, the vast cost of anticoccidial drugs and vaccines. In the poultry industry, conventional strategies for coccidiosis control mainly rely on chemoprophylaxis. Chemoprophylaxis can prevent coccidiosis but subclinical infections were widespread and because of development of drug resistance, the vast time and labor consuming, the new drugs development fall behind the drug resistance. Many researchers work on selection of new coccidiostat. Maybe it is a shortcut to development new coccidiostat on the base of studying the action mode of drugs in use. Halofuginone hydrobromide has been used for several years since1930s but little is known about its mode of action and target of coccidium. For the reasons aforementioned, we carried series of experiments to research the action mechanism of halofuginone hydrobromide against Eimeria tenella.
1. The epidemical investigation of chicken coccidiosis in Eastern China
An epidemical investigation in Eastern China was carried out to provide more information for the poultry industry.429feces samples from chickens in Zhejiang, Jiangsu, Shanghai, Anhui, Shandong, Jiangxi provinces were investigated in March and April,2011. The oocysts were isolated and identified. It is showed that there were116infected samples most of which were multiplex infection. The average infection rate of samples from the six provences was about27.1%, which arranged from30.0%to43.3%in Jiangxi, Shanghai, Shandong, Jiangsu, higher than the mean ratio. After the single oocyst propagation and multiplex-PCR identification, it is demonstrated that the predominant species were Eimeira maximum, E. tenella, E. necatrix and E. acervulina.
2. The anticoccidial effect evaluation of halofuginone hydrobromide
An experiment was carried out to verify the anticoccidial efficacy of halofuginone, a broad spectrum anticoccidial drugs.14-day-old chickens were inoculated with Eimeria tenella orally and then medicated with1.5mg/kg,3mg/kg,6mg/kg halofuginone, respectively. It was demonstrated that the body weight gains in groups of1.5mg/kg and3mg/kg were significantly higher than chickens in the positive control group. As the increasement of halofuginone concentration from1.5to6mg/kg feed, no bloody feces and cecal lesions were observed, but both oocysts production and body weight gains decreased. The anticoccidial index (ACI) of3mg/kg halofuginone was195.45. It is obvious that halofuginone is a good coccidiostat.
3. The safety evaluation of halofuginone hydrobromide
In order to evaluate the safety of halofuginone, chickens were provided with halofuginone for3weeks at the3times dose of recommended and the body weight gains, bloody and seum component was measured. The results showed that there was no statistical difference about the parameters between each groups. No obvious unnormal syndrome was observed after autopsy and histological examination. It is proposed halofuginone was safe to use as a coccidiostat.
4. The study on the action mode of halofuginone hydrobromide in vivo
Halofuginone hydrobromide has been used as an anticoccidial reagent for decades but very little is known about its mode of action. In this study, chickens were inoculated with Eimeria tenella oocysts and medicated with halofuginone at day0,1,2,3,4,5,6post inoculations (groups0,1,2,3,4,5,6, respectively). Chickens in group7were taken as challenge-unmedicated control and in group8 unchallenged-unmedicated control. The survival rate, body weight gains (BWG), oocysts production, cecal scores, bloody diarrhea and histological examinations were analyzed to evaluate the anticoccidial efficacy of halofuginone and to initially elucidate its mechanisms. Results showed that halofuginone can significantly enhance the BWG, and decrease both the oocyst shedding and cecal destruction caused by E. tenella infection. The histological slide examination noted that halofuginone was effective when provided0-2day post inoculation but only partially effective when applied3-7days post infection. The2nd generation schizonts treated with halofuginone appeared vacuolated and degenerated. It is concluded that halofuginone can inhibit the parasite's invasion of host cecal hypothetical cell at the early stages of life cycle and later disturb the parasite's development by vacuolation of the schizonts. The resulting abnormal schizonts could not divide into schizoites and were eventually eliminated by the host's immune response.
5. The study on the action mode of halofuginone hydrobromide in vitro
In order to investigate the action mode of halofuginone in vitro, the fresh unsporulated oocysts were inoculated with halofuginone hydrobromide. The sporulation rate, the changes of appearance and ultrastructures of oocysts, the relative levels of key genes expression and cell infective ability of sporozoites in vitro were evaluated. The results showed that halofuginone can inhibit spoogony of Eimeria tenella. After the treatment of halofuginone, the excystation of sporocysts was reduced, the appearance and ultrastructure of oocysts changed, and the cell infection was blocked. It is proposed that halofuginone can destroy the oocyst wall. According to the multi-genes expression analysis, the expression level of Eimeria tenella pyruvate kinase gene was down-regulated, and mannitol-1-phosphate dehydrogenase, microneme protein1, and heat shock protein90genes were up-regulated.
In conclusion, an epidemical survey of chicken coccidiosis in Eastern China was carried out and the Eimeria specieses was identified based on the morphology and multiplex-PCR assay; the experiments of anticoccidial efficacy and safety of halofuginone in chickens were carried out. The mode of action against Eimeria tenella was studied in vivo by restricted medication after infection. After that, we also studied the halofuginone impact on the sporogony and infectivity in vitro. At last, the target and action mechanism of halofuginone were illustrated which maybe helpful to poultry coccidiosis control and new coccidiostat development.
一、华东部分地区肉鸡球虫病的流行病学调查
为了了解华东地区肉鸡球虫病的流行情况,更好地为鸡球虫病的防治提供科学的依据,本项目组成员2011年3、4月份采集了来自华东地区的浙江省、江苏省、上海市、安徽省、山东省、江西省的养鸡场的429份鸡粪样品,进行了球虫卵囊的分离与鉴定。结果表明,有116份阳性样品,多数样品为两三种球虫的混合感染,六省市球虫病平均感染率为27.1%,江西、上海、山东、江苏四省市的球虫病感染率为30.0%-43.3%,高于平均水平。经单卵囊繁殖和多重PCR鉴定表明,上述六省市主要流行的球虫种类有四种,即巨型艾美耳球虫、柔嫩艾美耳球虫、毒害艾美耳球虫和堆型艾美耳球虫。
二、氢溴酸常山酮的抗球虫效果研究
氢溴酸常山酮是一类化学合成的高效广谱抗球虫药物,为了验证该产品对鸡球虫的作用效果,进行了本试验。本试验对14日龄鸡进行人工感染柔嫩艾美耳球虫,并用1.5mg/kg、3mg/kg和6mg/kg三种浓度的氢溴酸常山酮进行治疗。结果表明,与阳性对照组相比,1.5mg/kg、3mg/kg可以使球虫感染鸡的体重显著提高。随着药物浓度增加,没有观察到血便排出,没有观察到盲肠病变,但卵囊排出量和体重相对增重率都呈下降趋势。3mg/kg氢溴酸常山酮抗球虫指数高达195.45,表明该氢溴酸常山酮具有高效抗球虫效果。
三、氢溴酸常山酮的安全性评价
本实验以高于使用剂量1-3倍连续3周给药,通过鸡的增重情况、血液学与血液化学指标及组织病理学检查,评价了氢溴酸常山酮对鸡的安全性。实验结果表明,各实验组与对照组的血液化学指标和血液学指标无显著差异,尸体解剖未见各组织器官有明显的异常变化,组织学检查也未见有病理变化,说明氢溴酸常山酮对鸡只比较安全。
四、氢溴酸常山酮的体内作用机理研究
氢溴酸常山酮作为抗球虫药已有多年的使用历史,但是人们对其作用的机理却了解不多。为了研究其作用机理,本研究中分别对人工感染柔嫩艾美耳球虫后0、1、2、3、4、5、6天的鸡用3mg/kg饲料的氢溴酸常山酮进行治疗(相应的分别为第0、1、2、3、4、5、6组),观察不同给药时间的抗球虫效果。第7组的鸡只感染柔嫩艾美耳球虫而不治疗,作为感染对照组,第8组的鸡不感染不治疗,作为阴性对照组。感染后观察记录各组鸡的死亡率、体重增重率、卵囊下降率和盲肠病变记分,并且采集盲肠组织制备组织切片,综合评价氢溴酸常山酮的作用效果,初步探索其作用机理。实验结果表明,感染的当天或者感染后2天之内用氢溴酸常山酮治疗是完全有效的,如果感染2天后再用氢溴酸常山酮治疗只能起到部分保护效果或无保护效果。盲肠的组织切片观察表明,氢溴酸常山酮可以抑制柔嫩艾美耳球虫子孢子的入侵,氢溴酸常山酮治疗组的第二代裂殖体表现出空泡化和降解。因此,氢溴酸常山酮主要作用于球虫生活史的早期,阻止其入侵宿主细胞,然后通过抑制裂殖体分裂为裂殖子,使裂殖体空泡化达到破坏其发育的作用效果,导致裂殖体和裂殖子发育被抑制,发育速度减慢,从而被机体的免疫系统清除。
五、氢溴酸常山酮的体外作用机理研究
为了探索氢溴酸常山酮的体外作用机理,本实验对获得的新鲜的未孢子化柔嫩艾美耳球虫卵囊通过孢子化不同时间后给予氢溴酸常山酮孵育处理,再继续进行孢子化,观察了氢溴酸常山酮在体外条件下对孢子化的影响、卵囊的外观形态和超微结构的变化及部分有代表性关键基因的表达情况,并研究了子孢子体外处理后对细胞感染能力的影响。结果表明,氢溴酸常山酮可以抑制孢子生殖,使卵囊孢子化率降低,并且药物处理后的卵囊脱孢子囊能力下降,卵囊表面形态和细胞质及其超微结构等都发生明显异常,而且子孢子与氢溴酸常山酮体外条件下共孵育后对细胞的感染能力下降。根据结果推测,氢溴酸常山酮可能对卵囊壁的结构具有破坏作用。对多个基因的表达情况分析表明,氢溴酸常山酮作用后,丙醇酸激酶基因的表达下调,而1-磷酸甘露醇脱氢酶、微线体蛋白1和热激蛋白90基因表达上调。
综上所述,本研究对华东地区部分省市的鸡球虫病进行了流行病学调查,并用形态学方法和多重PCR方法对分离到的卵囊进行了虫种的鉴定；对氢溴酸常山酮进行了抗球虫效果实验及对鸡的安全性实验,并通过氢溴酸常山酮在球虫感染鸡体后不同时间给药探讨了抗柔嫩艾美耳球虫的作用机理,还在体外条件下,研究了氢溴酸常山酮对孢子生殖的影响,初步阐述了氢溴酸常山酮潜在的作用靶点及作用机理,但这些初步的结论还需要进一步的深入研究进行验证,这可能为球虫病的防控及药物开发提供借鉴意义。
Coccidiosis is caused by the intracellular protozoal Eimeria genus, which parasite in chicken's intestinal tract. Coccidiosis causes huge impaire to the poultry industry about$30billion annually because of the high mortality, reduced feed convertion rate, the vast cost of anticoccidial drugs and vaccines. In the poultry industry, conventional strategies for coccidiosis control mainly rely on chemoprophylaxis. Chemoprophylaxis can prevent coccidiosis but subclinical infections were widespread and because of development of drug resistance, the vast time and labor consuming, the new drugs development fall behind the drug resistance. Many researchers work on selection of new coccidiostat. Maybe it is a shortcut to development new coccidiostat on the base of studying the action mode of drugs in use. Halofuginone hydrobromide has been used for several years since1930s but little is known about its mode of action and target of coccidium. For the reasons aforementioned, we carried series of experiments to research the action mechanism of halofuginone hydrobromide against Eimeria tenella.
1. The epidemical investigation of chicken coccidiosis in Eastern China
An epidemical investigation in Eastern China was carried out to provide more information for the poultry industry.429feces samples from chickens in Zhejiang, Jiangsu, Shanghai, Anhui, Shandong, Jiangxi provinces were investigated in March and April,2011. The oocysts were isolated and identified. It is showed that there were116infected samples most of which were multiplex infection. The average infection rate of samples from the six provences was about27.1%, which arranged from30.0%to43.3%in Jiangxi, Shanghai, Shandong, Jiangsu, higher than the mean ratio. After the single oocyst propagation and multiplex-PCR identification, it is demonstrated that the predominant species were Eimeira maximum, E. tenella, E. necatrix and E. acervulina.
2. The anticoccidial effect evaluation of halofuginone hydrobromide
An experiment was carried out to verify the anticoccidial efficacy of halofuginone, a broad spectrum anticoccidial drugs.14-day-old chickens were inoculated with Eimeria tenella orally and then medicated with1.5mg/kg,3mg/kg,6mg/kg halofuginone, respectively. It was demonstrated that the body weight gains in groups of1.5mg/kg and3mg/kg were significantly higher than chickens in the positive control group. As the increasement of halofuginone concentration from1.5to6mg/kg feed, no bloody feces and cecal lesions were observed, but both oocysts production and body weight gains decreased. The anticoccidial index (ACI) of3mg/kg halofuginone was195.45. It is obvious that halofuginone is a good coccidiostat.
3. The safety evaluation of halofuginone hydrobromide
In order to evaluate the safety of halofuginone, chickens were provided with halofuginone for3weeks at the3times dose of recommended and the body weight gains, bloody and seum component was measured. The results showed that there was no statistical difference about the parameters between each groups. No obvious unnormal syndrome was observed after autopsy and histological examination. It is proposed halofuginone was safe to use as a coccidiostat.
4. The study on the action mode of halofuginone hydrobromide in vivo
Halofuginone hydrobromide has been used as an anticoccidial reagent for decades but very little is known about its mode of action. In this study, chickens were inoculated with Eimeria tenella oocysts and medicated with halofuginone at day0,1,2,3,4,5,6post inoculations (groups0,1,2,3,4,5,6, respectively). Chickens in group7were taken as challenge-unmedicated control and in group8 unchallenged-unmedicated control. The survival rate, body weight gains (BWG), oocysts production, cecal scores, bloody diarrhea and histological examinations were analyzed to evaluate the anticoccidial efficacy of halofuginone and to initially elucidate its mechanisms. Results showed that halofuginone can significantly enhance the BWG, and decrease both the oocyst shedding and cecal destruction caused by E. tenella infection. The histological slide examination noted that halofuginone was effective when provided0-2day post inoculation but only partially effective when applied3-7days post infection. The2nd generation schizonts treated with halofuginone appeared vacuolated and degenerated. It is concluded that halofuginone can inhibit the parasite's invasion of host cecal hypothetical cell at the early stages of life cycle and later disturb the parasite's development by vacuolation of the schizonts. The resulting abnormal schizonts could not divide into schizoites and were eventually eliminated by the host's immune response.
5. The study on the action mode of halofuginone hydrobromide in vitro
In order to investigate the action mode of halofuginone in vitro, the fresh unsporulated oocysts were inoculated with halofuginone hydrobromide. The sporulation rate, the changes of appearance and ultrastructures of oocysts, the relative levels of key genes expression and cell infective ability of sporozoites in vitro were evaluated. The results showed that halofuginone can inhibit spoogony of Eimeria tenella. After the treatment of halofuginone, the excystation of sporocysts was reduced, the appearance and ultrastructure of oocysts changed, and the cell infection was blocked. It is proposed that halofuginone can destroy the oocyst wall. According to the multi-genes expression analysis, the expression level of Eimeria tenella pyruvate kinase gene was down-regulated, and mannitol-1-phosphate dehydrogenase, microneme protein1, and heat shock protein90genes were up-regulated.
In conclusion, an epidemical survey of chicken coccidiosis in Eastern China was carried out and the Eimeria specieses was identified based on the morphology and multiplex-PCR assay; the experiments of anticoccidial efficacy and safety of halofuginone in chickens were carried out. The mode of action against Eimeria tenella was studied in vivo by restricted medication after infection. After that, we also studied the halofuginone impact on the sporogony and infectivity in vitro. At last, the target and action mechanism of halofuginone were illustrated which maybe helpful to poultry coccidiosis control and new coccidiostat development.
引文
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