福美双诱发肉鸡胫骨软骨发育不良差异表达基因的研究
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摘要
肉鸡胫骨软骨发育不良(Tibial Dyschondroplasia,TD)是肉鸡常见的骨骼疾病之一,以骺板成熟面形成不能血管化亦不能矿化的软骨栓为特点。TD的发生与品种、日粮的Ca/P比和阴阳离子的比例等因素有关,但TD的发病机理仍然不甚明了。目前,肉鸡胫骨软骨发育不良、肉鸡腹水综合征和肉鸡猝死综合征是公认的严重影响现代肉鸡养殖业的三大营养代谢病,每年给世界肉鸡养殖业造成巨大的经济损失。因此,肉鸡TD的发病机理和防治研究是目前世界上禽病研究的热点之一。
采用原位杂交、半定量RT-PCR等对肉鸡TD发生的分子机制研究,发现肉鸡息TD时,病变的软骨细胞生长分化相关基因存在表达紊乱、表达水平降低或表达缺失的现象。虽然采用以上方法对TD的发生的分子机理作了一定的阐明,但这些方法存在一定的盲目性和局限性,有必要采用新的研究手段对肉鸡TD发病的分子机理进行深入探讨,为抗TD肉鸡品种选育,治疗药物分子设计提供理论依据。
为此,本研究采用福美双诱发肉鸡TD,观察其动态组织病理学变化特征,分析其与自然发生TD的异同。同时,采用抑制性消减杂交(suppression substractivehvbridization,SSH)筛选鉴定与肉鸡TD相关的差异表达基因,并对差异基因表达的时序性进行了初步研究,取得了以下结果:
1福美双对肉鸡生长性能和胫骨软骨发育不良的影响
饲料中添加福美双可显著提高肉鸡TD发病率,达到94.44%。病理剖检结果症状典型,与自然发生的TD症状相似,且表现更为突出,胫骨干骺端粗大,胫骨弯曲;骺端生长板增厚,不钙化、无血管的软骨栓占据整个干骺端;而且与自然发病相比,还有福美双诱发TD独特的病理特征,即生长板质地松软,切面不完整,易断层。肉鸡生长受阻,体增重显著下降,料重比增加,严重影响了肉鸡生长性能。表明本研究成功建立了肉鸡TD模型。
2福美双诱发肉鸡胫骨软骨发育不良的动态组织病理学变化观察
动态组织病理学观察结果表明,福美双可促进软骨细胞极剧增殖,影响正常的细胞分化,导致细胞大量死亡,挤压血管,血管坏死,基质合成受阻,延缓软骨钙化,最终严重扰乱软骨成骨的调控机能;同时,将后期自然发生的TD和福美双诱发TD进行比较分析,发现二者组织病理学变化极为相似,从而进一步从细胞水平验证福美双成功诱发典型的肉鸡胫骨软骨发育不良,为深入研究TD的发生发展,以及研究预防TD的药物提供新的思路和重要线索。
3福美双诱发肉鸡胫骨软骨发育不良差异表达基因的筛选与鉴定
利用SSH技术构建了肉鸡胫骨软骨发育不良正反向差减cDNA文库。通过PCR和斑点杂交,从文库中筛选出40个差异表达EST。对差异表达EST进行序列分析,获得10个差异表达基因。以半定量RT-PCR对其中的7个差异表达基因进行了鉴定。试验结果表明,在TD发生中,Ⅹ型胶原(ColⅩ)、Ⅰ型胶原a1(ColⅠa1)、热休克蛋白(Hsp90)、NADH脱氢酶(NADH DH)等4个基因表达被上调;细胞色素c氧化酶Ⅲ(COXⅢ)、Matrilin-3(MATN3)、碳酸酐酶(CA2)等3个基因表达被下调。ColⅩ通过形成基质小泡使钙离子流入并区划基质成分来调节软骨内骨化;ColⅠ脯氨酰的羟化作用是骨形成的关键;CA2参与骨骼的发育与重构;Matrilin-3作为一种表面成分,通过与ColⅨ直接相连,或者以COMP为接头间接地与ColⅨ相连,可与基质大分子网络互联并且调节纤维和纤维基质间(蛋白聚糖)的相互作用,有助于骨化时基质的成熟;Hsp90具有多种细胞功能,作为分子伴侣调节蛋白构象、参与蛋白组装、参与生物信号转导、抑制细胞凋亡、延缓衰老、抗应激等;NADH DH和COXⅢ参与细胞的电子传递,与细胞的能量代谢有关。对TD中干扰软骨细胞分化的差异表达基因的鉴定,有助于TD分子机理的研究,从而为遗传选育消除机能紊乱开辟了新的途径。
4福美双诱发肉鸡胫骨软骨发育不良基因差异表达的时序性研究
用SSH技术所筛选的上调表达基因Ⅹ型胶原(ColⅩ)、Ⅰ型胶原(ColⅠ)、热休克蛋白(Hsp90)和下调表达基因Chondromodulin-1(ChM-1)、碳酸酐酶(CA2)、烯醇酶1(ENO1)经原位杂交验证均为差异表达基因,这对于深入研究TD的发病机理奠定了理论基础。
通过对Ⅰ型胶原(ColⅠ)、热休克蛋白(Hsp90)免疫组化时序性研究显示,对于上调基因其细胞中相应蛋白也随之增加,表明TD发生可能主要是引起基因的表达紊乱所致,从而为TD分子致病机理的研究明确了新的方向。
Tibial dyschondroplasia(TD),characteristic of accumulated growth plate cartilage into metaphyseal region of the tibiotarsus,bone deformity and lameness is one of diseases that affect fast-growing birds worldwide.The development of TD is involved in such factors as genotype,the calcium/phosphorus ratio and acid-base balance of the diet. Although many factors that cause TD have been identified,the detailed etiology of TD is not fully understood.TD along with ascite and sudden death are regarded as the three nutritional and metabolic diseases that severely affect broiler industry.As occurrence of TD leads to big loss to broiler industry worldwide,close attentions have been paid to identify its etiolgy.
Some molecular techniques,such as in situ hybridization and semi-quantitative RT-PCR have been used to investigate the molecular mechanism of this disease.The results showed that expression of some genes were disordered,or its levels were lowered, or it was absent during the development of TD.As these techniques have limitations and blindness,new methods must be used to investigate the molecular mechanisms of TD in broilers,which will provide theoretical basis to breed TD resilient broiler species and to develop medicines for treatment of TD.
In this study,TD in broilers was induced by using thiram and the dynamic pathological changes were observed and compared with those occurred spontaneously in TD.Furthermore,suppression substractive hybridization was employed to identify the differentially expressed genes during the development of TD.Gene differential expression at different stages in the development of TD was validated by using in situ hybridization and immunohistochemistry.The results were as follows:
1 Effects of thiram on growth performance and tibial dyschondroplasis in Broiler Chickens
Thiram significantly increased incidence of TD(94.44%).The pathologic symptoms in the induced TD were similar to those of spontaneous TD,but were more severe than those of spontaneous TD.The lesion composed of noncalcified,nonvascularized, accumulated cartilage that can extend from the epiphyseal growth plate to the all metaphysic.The distinct character was that of the fragile growth plate.Broiler growth was slow,body weight was decreased,feed to gain ratio(F/G) was increased,and growth performance was affected.The model of thiram inducing TD may be regarded as an ideal animal model to study the mechanisms of occurrence and the progression of this disease.
2 Dynamic changes of histopathology of tibial dyschondroplasis in broiler chickens induced by thiram
The study of typical pathohistological changes in the metaphyseal region of the tibiotarsus in experimental broilers fed diet supplemented with thiram showed that the thiram promoted chondrocyte proliferation,resulted in death of large number of cells, compressed blood vessels which led to necrosis of blood vessel,hindered synthesis of extracellular matrix,deferred endochondral calcification.As a result,it severely disturbed the functioning of regulating endochondral bone.By comparison of spontaneously occurring TD and that induced by thiram,the result demonstrated that thiram could induce tibial dyschondroplasia,led to the changes in morphology and histology of growth plates,which were similar to those of spontaneously occurring TD.Therefore the study of dynamic changes may provide important thinking and clue to studying the molecular mechanism of TD development,and to screening for the effects of different micronutrients against the pathogenesis of tibial dyschondroplasia.
3 Selection and identification of differentially expressed genes of tibial dyschondroplasa in broiler chikens induced by thiram
To investigate the molecular mechanisms of broiler tibial dyschondroplasia,forward and reverse subtracted cDNA libraries were built with cDNA from tibial cartilage growth plate of thiram-treated and normal broiler chickens by suppression subtractive hybridization(SSH) technology.Three hundred and fifty-six available clones were selected randomly in the two libraries.40 differentially expressed EST were selected by means of PCR and reverse dot blotting.The sequencing results revealed that these genes represented 10 known genes using blasting on-line with the GenBank dbase.Seven genes were differentially expressed in TD and normal broilers,which were revealed by semi-quantitative RT-PCR.TypeⅩcollagen(ColⅩ),TypeⅠcollagen(ColⅠ),heat shock protein 90(Hsp90) and NADH dehydrogenase(NADH DH) gene expressing levels was up-regulated.Cytochrome C oxidaseⅢ(COXⅢ),Matrilin-3(MATN3) and carbonic anhydraseⅡ(CA2) gene was down-regulated.TypeⅩcollagen facilitates endochondral ossification by forming matrix vesicles,Ca~(2+) influxing into matrix vesicles and compartmentalizing matrix components.Eprolyl 3-hydroxylation of typeⅠcollagen is important for bone form.CA2 is involved in skeleton development and reconstitution. Matrilin-3 can be seen as an interface component,capable of interconnecting macromolecular networks,mediating interactions between cartilage fibrils and the extrafibrillar matrix and contributing to matrical maturation by a direct interaction with collagenⅨand indirectly with COMP serving as an adapter.As molecular chaperones, Hsp90 had such functions as regulating protein conformation,participating protein assemble and install,involving in signal transduction,inhibiting apoptosis,delaying aging and resisting stress.NADH DH and COXⅢrelated to cell metabolism by transferring electron.This identification of these differentially expressed genes involved in the perturbation of chondrocyte differentiation in TD is significant for studying the molecular mechanisms of TD and may open the new way to the elimination of the disorder by genetic selection.
4 Gene differential expression at different stages in the development of tibial dyschondroplasa induced by thiram in Broiler chikens
The up-regulated genes typeⅩcollogen(ColⅩ),typeⅠcollogen(ColⅠ),heat shock protein 90(Hsp90) and down-regulated genes Chondromodulin-1(ChM-1),carbonic anhydraseⅡ(CA2),enolase1(ENO1) were revealed by in situ hybridization.
The results demonstrated that TD chondrocytes synthesized elevated levels of typeⅠcollagen and heat shock protein 90 mRNA by in situ hybridization.Protein levels of TypeⅠcollagen and heat shock protein 90 were also increased revealed by immunohistochemistry.So it was suggested that the perturbation of genes expression led to TD,as a result,the research provides a new direction for studying the molecular mechanisms of TD.
采用原位杂交、半定量RT-PCR等对肉鸡TD发生的分子机制研究,发现肉鸡息TD时,病变的软骨细胞生长分化相关基因存在表达紊乱、表达水平降低或表达缺失的现象。虽然采用以上方法对TD的发生的分子机理作了一定的阐明,但这些方法存在一定的盲目性和局限性,有必要采用新的研究手段对肉鸡TD发病的分子机理进行深入探讨,为抗TD肉鸡品种选育,治疗药物分子设计提供理论依据。
为此,本研究采用福美双诱发肉鸡TD,观察其动态组织病理学变化特征,分析其与自然发生TD的异同。同时,采用抑制性消减杂交(suppression substractivehvbridization,SSH)筛选鉴定与肉鸡TD相关的差异表达基因,并对差异基因表达的时序性进行了初步研究,取得了以下结果:
1福美双对肉鸡生长性能和胫骨软骨发育不良的影响
饲料中添加福美双可显著提高肉鸡TD发病率,达到94.44%。病理剖检结果症状典型,与自然发生的TD症状相似,且表现更为突出,胫骨干骺端粗大,胫骨弯曲;骺端生长板增厚,不钙化、无血管的软骨栓占据整个干骺端;而且与自然发病相比,还有福美双诱发TD独特的病理特征,即生长板质地松软,切面不完整,易断层。肉鸡生长受阻,体增重显著下降,料重比增加,严重影响了肉鸡生长性能。表明本研究成功建立了肉鸡TD模型。
2福美双诱发肉鸡胫骨软骨发育不良的动态组织病理学变化观察
动态组织病理学观察结果表明,福美双可促进软骨细胞极剧增殖,影响正常的细胞分化,导致细胞大量死亡,挤压血管,血管坏死,基质合成受阻,延缓软骨钙化,最终严重扰乱软骨成骨的调控机能;同时,将后期自然发生的TD和福美双诱发TD进行比较分析,发现二者组织病理学变化极为相似,从而进一步从细胞水平验证福美双成功诱发典型的肉鸡胫骨软骨发育不良,为深入研究TD的发生发展,以及研究预防TD的药物提供新的思路和重要线索。
3福美双诱发肉鸡胫骨软骨发育不良差异表达基因的筛选与鉴定
利用SSH技术构建了肉鸡胫骨软骨发育不良正反向差减cDNA文库。通过PCR和斑点杂交,从文库中筛选出40个差异表达EST。对差异表达EST进行序列分析,获得10个差异表达基因。以半定量RT-PCR对其中的7个差异表达基因进行了鉴定。试验结果表明,在TD发生中,Ⅹ型胶原(ColⅩ)、Ⅰ型胶原a1(ColⅠa1)、热休克蛋白(Hsp90)、NADH脱氢酶(NADH DH)等4个基因表达被上调;细胞色素c氧化酶Ⅲ(COXⅢ)、Matrilin-3(MATN3)、碳酸酐酶(CA2)等3个基因表达被下调。ColⅩ通过形成基质小泡使钙离子流入并区划基质成分来调节软骨内骨化;ColⅠ脯氨酰的羟化作用是骨形成的关键;CA2参与骨骼的发育与重构;Matrilin-3作为一种表面成分,通过与ColⅨ直接相连,或者以COMP为接头间接地与ColⅨ相连,可与基质大分子网络互联并且调节纤维和纤维基质间(蛋白聚糖)的相互作用,有助于骨化时基质的成熟;Hsp90具有多种细胞功能,作为分子伴侣调节蛋白构象、参与蛋白组装、参与生物信号转导、抑制细胞凋亡、延缓衰老、抗应激等;NADH DH和COXⅢ参与细胞的电子传递,与细胞的能量代谢有关。对TD中干扰软骨细胞分化的差异表达基因的鉴定,有助于TD分子机理的研究,从而为遗传选育消除机能紊乱开辟了新的途径。
4福美双诱发肉鸡胫骨软骨发育不良基因差异表达的时序性研究
用SSH技术所筛选的上调表达基因Ⅹ型胶原(ColⅩ)、Ⅰ型胶原(ColⅠ)、热休克蛋白(Hsp90)和下调表达基因Chondromodulin-1(ChM-1)、碳酸酐酶(CA2)、烯醇酶1(ENO1)经原位杂交验证均为差异表达基因,这对于深入研究TD的发病机理奠定了理论基础。
通过对Ⅰ型胶原(ColⅠ)、热休克蛋白(Hsp90)免疫组化时序性研究显示,对于上调基因其细胞中相应蛋白也随之增加,表明TD发生可能主要是引起基因的表达紊乱所致,从而为TD分子致病机理的研究明确了新的方向。
Tibial dyschondroplasia(TD),characteristic of accumulated growth plate cartilage into metaphyseal region of the tibiotarsus,bone deformity and lameness is one of diseases that affect fast-growing birds worldwide.The development of TD is involved in such factors as genotype,the calcium/phosphorus ratio and acid-base balance of the diet. Although many factors that cause TD have been identified,the detailed etiology of TD is not fully understood.TD along with ascite and sudden death are regarded as the three nutritional and metabolic diseases that severely affect broiler industry.As occurrence of TD leads to big loss to broiler industry worldwide,close attentions have been paid to identify its etiolgy.
Some molecular techniques,such as in situ hybridization and semi-quantitative RT-PCR have been used to investigate the molecular mechanism of this disease.The results showed that expression of some genes were disordered,or its levels were lowered, or it was absent during the development of TD.As these techniques have limitations and blindness,new methods must be used to investigate the molecular mechanisms of TD in broilers,which will provide theoretical basis to breed TD resilient broiler species and to develop medicines for treatment of TD.
In this study,TD in broilers was induced by using thiram and the dynamic pathological changes were observed and compared with those occurred spontaneously in TD.Furthermore,suppression substractive hybridization was employed to identify the differentially expressed genes during the development of TD.Gene differential expression at different stages in the development of TD was validated by using in situ hybridization and immunohistochemistry.The results were as follows:
1 Effects of thiram on growth performance and tibial dyschondroplasis in Broiler Chickens
Thiram significantly increased incidence of TD(94.44%).The pathologic symptoms in the induced TD were similar to those of spontaneous TD,but were more severe than those of spontaneous TD.The lesion composed of noncalcified,nonvascularized, accumulated cartilage that can extend from the epiphyseal growth plate to the all metaphysic.The distinct character was that of the fragile growth plate.Broiler growth was slow,body weight was decreased,feed to gain ratio(F/G) was increased,and growth performance was affected.The model of thiram inducing TD may be regarded as an ideal animal model to study the mechanisms of occurrence and the progression of this disease.
2 Dynamic changes of histopathology of tibial dyschondroplasis in broiler chickens induced by thiram
The study of typical pathohistological changes in the metaphyseal region of the tibiotarsus in experimental broilers fed diet supplemented with thiram showed that the thiram promoted chondrocyte proliferation,resulted in death of large number of cells, compressed blood vessels which led to necrosis of blood vessel,hindered synthesis of extracellular matrix,deferred endochondral calcification.As a result,it severely disturbed the functioning of regulating endochondral bone.By comparison of spontaneously occurring TD and that induced by thiram,the result demonstrated that thiram could induce tibial dyschondroplasia,led to the changes in morphology and histology of growth plates,which were similar to those of spontaneously occurring TD.Therefore the study of dynamic changes may provide important thinking and clue to studying the molecular mechanism of TD development,and to screening for the effects of different micronutrients against the pathogenesis of tibial dyschondroplasia.
3 Selection and identification of differentially expressed genes of tibial dyschondroplasa in broiler chikens induced by thiram
To investigate the molecular mechanisms of broiler tibial dyschondroplasia,forward and reverse subtracted cDNA libraries were built with cDNA from tibial cartilage growth plate of thiram-treated and normal broiler chickens by suppression subtractive hybridization(SSH) technology.Three hundred and fifty-six available clones were selected randomly in the two libraries.40 differentially expressed EST were selected by means of PCR and reverse dot blotting.The sequencing results revealed that these genes represented 10 known genes using blasting on-line with the GenBank dbase.Seven genes were differentially expressed in TD and normal broilers,which were revealed by semi-quantitative RT-PCR.TypeⅩcollagen(ColⅩ),TypeⅠcollagen(ColⅠ),heat shock protein 90(Hsp90) and NADH dehydrogenase(NADH DH) gene expressing levels was up-regulated.Cytochrome C oxidaseⅢ(COXⅢ),Matrilin-3(MATN3) and carbonic anhydraseⅡ(CA2) gene was down-regulated.TypeⅩcollagen facilitates endochondral ossification by forming matrix vesicles,Ca~(2+) influxing into matrix vesicles and compartmentalizing matrix components.Eprolyl 3-hydroxylation of typeⅠcollagen is important for bone form.CA2 is involved in skeleton development and reconstitution. Matrilin-3 can be seen as an interface component,capable of interconnecting macromolecular networks,mediating interactions between cartilage fibrils and the extrafibrillar matrix and contributing to matrical maturation by a direct interaction with collagenⅨand indirectly with COMP serving as an adapter.As molecular chaperones, Hsp90 had such functions as regulating protein conformation,participating protein assemble and install,involving in signal transduction,inhibiting apoptosis,delaying aging and resisting stress.NADH DH and COXⅢrelated to cell metabolism by transferring electron.This identification of these differentially expressed genes involved in the perturbation of chondrocyte differentiation in TD is significant for studying the molecular mechanisms of TD and may open the new way to the elimination of the disorder by genetic selection.
4 Gene differential expression at different stages in the development of tibial dyschondroplasa induced by thiram in Broiler chikens
The up-regulated genes typeⅩcollogen(ColⅩ),typeⅠcollogen(ColⅠ),heat shock protein 90(Hsp90) and down-regulated genes Chondromodulin-1(ChM-1),carbonic anhydraseⅡ(CA2),enolase1(ENO1) were revealed by in situ hybridization.
The results demonstrated that TD chondrocytes synthesized elevated levels of typeⅠcollagen and heat shock protein 90 mRNA by in situ hybridization.Protein levels of TypeⅠcollagen and heat shock protein 90 were also increased revealed by immunohistochemistry.So it was suggested that the perturbation of genes expression led to TD,as a result,the research provides a new direction for studying the molecular mechanisms of TD.
引文
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