日粮硒对雏鸡免疫功能影响的机理研究
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摘要
本研究以病理学、免疫学和流式细胞术等主要方法,研究高硒和低硒日粮对雏鸡免疫器官生长发育、免疫器官细胞增殖与凋亡、脾脏抗氧化能力、以及细胞免疫、体液免疫和红细胞免疫功能的动态影响规律和机理,为基础研究和生产实践提供理论依据。研究包括两个动物试验。
实验一日粮高硒对雏鸡免疫功能影响的机理研究
选用1日龄艾维茵肉鸡300只,随机分成5组,分别喂以玉米豆粕型对照日粮(Se 0.2mg/kg)和高硒日粮(Se 1mg/kg高硒Ⅰ组,Se 5mg/kg高硒Ⅱ组,Se 10mg/kg高硒Ⅲ组,Se 15mg/kg高硒Ⅳ组),试验期6周。结果显示:①高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊重量减轻,脏器指数减小,器官生长发育受阻。眼观变化见胸腺、脾脏和法氏囊体积缩小。组织学观察显示,高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺皮质和髓质区淋巴细胞数量减少,胸腺小体体积增大,中央区细胞核碎片增多;脾脏和法氏囊淋巴细胞减少。胸腺、脾脏和法氏囊的超微病理变化以淋巴细胞和网状细胞的线粒体病变和凋亡细胞数量增多为特征。②流式细胞术检测表明,高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊细胞静止期(G_0G_1)升高,合成期(S)、分裂(G_2M)期下降,细胞分裂指数(PI)下降。日粮硒含量越高,变化趋势越明显,且与脏器指数的下降趋势一致。③Annexin V-FITC染色检测结果显示,高硒Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊的凋亡百分率明显升高。TUNEL染色的定性研究显示,高硒Ⅲ、Ⅳ组的胸腺、脾脏和法氏囊中,染成棕黄色的凋亡细胞较对照组增多。电镜观察,凋亡细胞见典型的超微结构特征,细胞核染色质浓缩边移,呈马蹄形或新月形。凋亡细胞增多的改变与免疫器官生长发育受阻、淋巴细胞减少的变化一致。④高硒Ⅱ、Ⅲ、Ⅳ组雏鸡外周血CD3+T细胞、CD3+CD4+T细胞和CD3+CD8+T细胞百分率程度不同地降低;高硒Ⅲ、Ⅳ组雏鸡外周血淋巴细胞对刀豆素A刺激的细胞增殖反应减弱;血清IL-2含量降低。雏鸡细胞免疫功能受损。⑤高硒Ⅱ、Ⅲ、Ⅳ组雏鸡血清IgG含量下降;高硒Ⅳ组雏鸡血清IgA和IgM含量。高硒Ⅲ、Ⅳ组血清总蛋白、白蛋白和球蛋白含量下降。雏鸡体液免疫功能受损。⑥高硒Ⅱ、Ⅲ、IV组红细胞C_3bR花环率降低,免疫复合物IC花环率升高,红细胞免疫粘附功能受损。⑦高硒Ⅲ、Ⅳ组脾硒含量升高,脾脏GSH-Px、SOD酶和CAT酶活性下降、总抗氧化能力降低、MDA蓄积,造成脾脏结构和功能损伤,细胞免疫和体液免疫功能受损。
实验二日粮低硒对雏鸡免疫功能影响的机理研究
选用1日龄艾维茵肉鸡120只,随机分成2组,硒缺乏组日粮由购自西昌冕宁的玉米、黄豆和小麦配制而成,硒含量为0.0342mg/kg,以亚硒酸钠为硒源使日粮硒水平达0.2mg/kg构成对照日粮,试验期6周。结果显示:①低硒组雏鸡胸腺、脾脏和法氏囊重量减轻,脏器指数减小。眼观变化见胸腺、脾脏和法氏囊体积小于对照组。组织学观察低硒组胸腺、脾脏和法氏囊淋巴细胞减少,其中法氏囊在42日龄时,淋巴滤泡内的淋巴细胞几近耗竭,网状细胞增生。胸腺、脾脏和法氏囊的超微病理变化以淋巴细胞和网状细胞的线粒体病变和凋亡细胞数量增多为特征。②低硒组雏鸡胸腺、脾脏和法氏囊细胞G_0G_1期升高,G2M期和PI降低,细胞分裂增殖能力受抑,免疫器官生长发育受阻。③流式细胞术、TUNEL染色法和电镜观察显示,低硒组胸腺、脾脏和法氏囊的凋亡细胞增多,此改变与免疫器官淋巴细胞数量减少的病变密切相关。④低硒组雏鸡外周血CD3+T细胞和CD3+CD8+T细胞百分率程度不同地降低;外周血淋巴细胞对刀豆素A刺激的细胞增殖反应减弱;血清IL-2含量降低。雏鸡细胞免疫功能受损。⑤低硒组雏鸡血清IgG、IgA和IgM含量降低,血清总蛋白、白蛋白和球蛋白含量降低,体液免疫功能受损。⑥低硒组雏鸡红细胞C_3bR花环率,IC花环率变化不明显,发生原发性免疫功能低下。⑦低硒组脾硒含量降低,脾脏GSH-Px、CAT活性和总抗氧化能力降低,MDA含量增加,脾脏组织结构和功能受损。
上述研究结果表明:低硒(0.0342mg/kg)和5mg/kg及其以上高硒引起免疫器官淋巴细胞分裂增殖能力下降,凋亡细胞数量增多,脾脏组织中抗氧化酶活性降低、过氧化产物MDA蓄积,导致免疫器官生长发育不良,细胞免疫、体液免疫和红细胞免疫功能降低。细胞分裂增殖能力下降、凋亡细胞增多和抗氧化能力降低、过氧化产物蓄积是免疫功能受损的病理基础和机理。本研究首次全面系统研究了日粮硒对艾维茵肉鸡免疫器官发育和免疫功能的影响规律及机制,为养鸡生产上确定硒的合理添加剂量、诊断和防治硒缺乏症和硒中毒症提供了理论依据。
Two experiments were conducted to study on the effect of high selenium(Se) and low Se on the structure and function of immune system in chickens. The cell cycle and apoptosis of immune organs, the function of cellular, humoral and erythrocyte immunity, and the anti-oxidative function of spleen were determined by the means of pathology, immunology and flow cytometry.
Experiment 1 Mechanisms of the immune functional changes caused by high Se in chickens
300 one-day-old avian chickens were randomly divided into five groups, and fed on diets as follows: corn-bean control diet (Se 0.2mg/kg) and high Se diets (Se 1mg/kg, high Se group I ; Se 5mg/kg, high Se group II; Se 10mg/kg, high Se group III; Se 15mg/kg, high Se group IV) for 6 weeks. The results were as follow:①The volume, weight and relative weight of thymus, spleen and bursa of Fabricius(BF) were significantly decreased in high Se groups II , III and IV when compared with those of control group. Histopathologically, decreased lymphocytes, enlarged thymic corpuscle and increased nuclear debris in the center of corpuscles were appeared in the thymus. Decreased lymphocytes were observed in spleen and BF of high Se groups II, III and IV. Ultrastructurally, mitochondria injury and increased apoptotic cells with condensed nuclei were appeared in immune organs.②Cell cycle analysis by flow cytometry showed that the G_0G_1 phase was increased, and the S phase, G_2M phase and proliferation index(PI) were decreased in thymus, spleen and BF of high Se II, III and IV. Changes of cell cycle were progressed with the dietary Se level increasing, and consistent with the changes of organs relative weight.③The percentage of Annexin-V positive cells measured by flow cytometry was higher in high Se groups II, III and IV than in control group. TUNEL assay showed that there were increased frequencies of apoptotic cells in high Se groups III and IV. Characteristic of ultrastructural features on apoptotic cells was nuclear membrane breakdown, chromatin condensation with horseshoe-like or crescent shapes. The results were consistent with suppressed growth and decreased lymphocytes of immune organs.④The percentage of CD3+ T cells, CD3+CD4+ T cells and CD3+CD8+ T cells were decreased variably in high Se groups II, III and IV groups when compared with those of control group. The mitogenic reponse of T lymphocytes to ConA and the contents of serum IL-2 were decreased in high Se groups III and IV than in control group. The results suggested that the cellular immune function of chickens was impaired by high Se.⑤The contents of serum IgG were lower in high Se groups III and IV than in control group. The contents of serum IgA and IgM were decreased in high Se group IV. Contents of serum total protein, albumin and globulin were decreased variably in high Se groups III and IV. The results suggested that humoral immune function of chickens was impaired by high Se.⑥The ratio of IC rosette was increased and the ratio of C_3bR rosette decreased in High Se II, III and IV. Immune adherence function of erythrocyte was impaired by high Se.⑦The splenic Se and MDA contents were higher, and splenic GSH-Px, SOD and CAT activities, and total antioxidative ability were lower in high Se groups III and IV than in control group. The structure and function of spleen were impaired. The cellular and humoral function were decreased.
Experiment 2 Mechanisms of the immune functional impairment caused by low Se in chickens
120 one-day-old avian chickens were randomly divided into two groups. Corn, yellow bean and wheat, which were bought from Mianling count were used to formulated low Se diet (Se 0.0342 mg/kg). Sodium selenite was mixed into the low Se diet to produce 0.2 mg/kg Se diet (control group). Broilers were fed the diets for 6 weeks. The results were as follow:①The weight and relative weight of thymus, spleen and BF were significantly decreased in low Se group when compared with those of control group. Anatomically, the volume of immune organs were smaller in low Se group than in control group. Histopathologically, lymphocytes in thymus, spleen and BF were decreased. At 42 days of age, atrophy BF showed infolded epithelium, shrinked follicle with reticuloendothelial cells proliferation. Ultrastructurally, mitochondria injury and increased apoptotic cells with condensed nuclei were observed.②Cell cycle analysis by flow cytometry showed that the G_0G_1 phase was increased, and the G_2M phase and proliferation index(PI) were decreased in thymus, spleen and BF of low Se group. Decreased proliferative function induced growth retardation of immune organs.③Increased apoptotic cells in thymus, spleen and BF were observed by the methods of flow cytometry, TUNEL stain and electron microscope. The results were associated with hypocellular of immune organs.④The percentage of CD3+ T cells and CD3+CD8+ T cells were decreased in low Se group. The mitogenic reponse of T lymphocytes to ConA and the content of serum IL-2 were lower in low Se group than in control group. The results showed that the cellular immune function of chickens was impaired by low Se.⑤The content of serum IgG, IgA, IgM, total protein, albumin and globulin were decreased iIn low Se group. The humoral immune function of chickens were decreased.⑥The ratio of C_3bR rosette was decreased, and the ratio of IC rosette had no obvious change in low Se group. Primary lesion of erythrocyte's adherence function was induced by low Se.⑦The contents of splenic Se, total antioxidative ability, GSH-Px and CAT activities were decreased, and MDA contents in spleen increased in low Se group. The structure and function of spleen were impaired.
The aforementioned results showed that the proliferative function of immune organs and activities of antioxidase in spleen were decreased, and the percentage of apoptotic cells of imunne organs and splenic MDA contents were increased by low Se (0.0342 mg/kg) and high Se (more than 5 mg/kg), which induced growth retardation of immune organs and decrease of cellular immune, humoral immune and erythrocyte's adherence function. The mechanisms of immune injury were closely related to the decreased cellular proliferative function, increased cellular apoptosis, decreased antioxidative ability and accumulated peroxidative products in the immune organs. In the present study, the effect of dietary Se on the structure and function of immune system was systematically researched for the first time. The results provided theoretical basis for determining reasonable supplemental dose of Se and preventing Se-deficiency and selenosis in chickens.
实验一日粮高硒对雏鸡免疫功能影响的机理研究
选用1日龄艾维茵肉鸡300只,随机分成5组,分别喂以玉米豆粕型对照日粮(Se 0.2mg/kg)和高硒日粮(Se 1mg/kg高硒Ⅰ组,Se 5mg/kg高硒Ⅱ组,Se 10mg/kg高硒Ⅲ组,Se 15mg/kg高硒Ⅳ组),试验期6周。结果显示:①高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊重量减轻,脏器指数减小,器官生长发育受阻。眼观变化见胸腺、脾脏和法氏囊体积缩小。组织学观察显示,高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺皮质和髓质区淋巴细胞数量减少,胸腺小体体积增大,中央区细胞核碎片增多;脾脏和法氏囊淋巴细胞减少。胸腺、脾脏和法氏囊的超微病理变化以淋巴细胞和网状细胞的线粒体病变和凋亡细胞数量增多为特征。②流式细胞术检测表明,高硒Ⅱ、Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊细胞静止期(G_0G_1)升高,合成期(S)、分裂(G_2M)期下降,细胞分裂指数(PI)下降。日粮硒含量越高,变化趋势越明显,且与脏器指数的下降趋势一致。③Annexin V-FITC染色检测结果显示,高硒Ⅲ、Ⅳ组雏鸡胸腺、脾脏和法氏囊的凋亡百分率明显升高。TUNEL染色的定性研究显示,高硒Ⅲ、Ⅳ组的胸腺、脾脏和法氏囊中,染成棕黄色的凋亡细胞较对照组增多。电镜观察,凋亡细胞见典型的超微结构特征,细胞核染色质浓缩边移,呈马蹄形或新月形。凋亡细胞增多的改变与免疫器官生长发育受阻、淋巴细胞减少的变化一致。④高硒Ⅱ、Ⅲ、Ⅳ组雏鸡外周血CD3+T细胞、CD3+CD4+T细胞和CD3+CD8+T细胞百分率程度不同地降低;高硒Ⅲ、Ⅳ组雏鸡外周血淋巴细胞对刀豆素A刺激的细胞增殖反应减弱;血清IL-2含量降低。雏鸡细胞免疫功能受损。⑤高硒Ⅱ、Ⅲ、Ⅳ组雏鸡血清IgG含量下降;高硒Ⅳ组雏鸡血清IgA和IgM含量。高硒Ⅲ、Ⅳ组血清总蛋白、白蛋白和球蛋白含量下降。雏鸡体液免疫功能受损。⑥高硒Ⅱ、Ⅲ、IV组红细胞C_3bR花环率降低,免疫复合物IC花环率升高,红细胞免疫粘附功能受损。⑦高硒Ⅲ、Ⅳ组脾硒含量升高,脾脏GSH-Px、SOD酶和CAT酶活性下降、总抗氧化能力降低、MDA蓄积,造成脾脏结构和功能损伤,细胞免疫和体液免疫功能受损。
实验二日粮低硒对雏鸡免疫功能影响的机理研究
选用1日龄艾维茵肉鸡120只,随机分成2组,硒缺乏组日粮由购自西昌冕宁的玉米、黄豆和小麦配制而成,硒含量为0.0342mg/kg,以亚硒酸钠为硒源使日粮硒水平达0.2mg/kg构成对照日粮,试验期6周。结果显示:①低硒组雏鸡胸腺、脾脏和法氏囊重量减轻,脏器指数减小。眼观变化见胸腺、脾脏和法氏囊体积小于对照组。组织学观察低硒组胸腺、脾脏和法氏囊淋巴细胞减少,其中法氏囊在42日龄时,淋巴滤泡内的淋巴细胞几近耗竭,网状细胞增生。胸腺、脾脏和法氏囊的超微病理变化以淋巴细胞和网状细胞的线粒体病变和凋亡细胞数量增多为特征。②低硒组雏鸡胸腺、脾脏和法氏囊细胞G_0G_1期升高,G2M期和PI降低,细胞分裂增殖能力受抑,免疫器官生长发育受阻。③流式细胞术、TUNEL染色法和电镜观察显示,低硒组胸腺、脾脏和法氏囊的凋亡细胞增多,此改变与免疫器官淋巴细胞数量减少的病变密切相关。④低硒组雏鸡外周血CD3+T细胞和CD3+CD8+T细胞百分率程度不同地降低;外周血淋巴细胞对刀豆素A刺激的细胞增殖反应减弱;血清IL-2含量降低。雏鸡细胞免疫功能受损。⑤低硒组雏鸡血清IgG、IgA和IgM含量降低,血清总蛋白、白蛋白和球蛋白含量降低,体液免疫功能受损。⑥低硒组雏鸡红细胞C_3bR花环率,IC花环率变化不明显,发生原发性免疫功能低下。⑦低硒组脾硒含量降低,脾脏GSH-Px、CAT活性和总抗氧化能力降低,MDA含量增加,脾脏组织结构和功能受损。
上述研究结果表明:低硒(0.0342mg/kg)和5mg/kg及其以上高硒引起免疫器官淋巴细胞分裂增殖能力下降,凋亡细胞数量增多,脾脏组织中抗氧化酶活性降低、过氧化产物MDA蓄积,导致免疫器官生长发育不良,细胞免疫、体液免疫和红细胞免疫功能降低。细胞分裂增殖能力下降、凋亡细胞增多和抗氧化能力降低、过氧化产物蓄积是免疫功能受损的病理基础和机理。本研究首次全面系统研究了日粮硒对艾维茵肉鸡免疫器官发育和免疫功能的影响规律及机制,为养鸡生产上确定硒的合理添加剂量、诊断和防治硒缺乏症和硒中毒症提供了理论依据。
Two experiments were conducted to study on the effect of high selenium(Se) and low Se on the structure and function of immune system in chickens. The cell cycle and apoptosis of immune organs, the function of cellular, humoral and erythrocyte immunity, and the anti-oxidative function of spleen were determined by the means of pathology, immunology and flow cytometry.
Experiment 1 Mechanisms of the immune functional changes caused by high Se in chickens
300 one-day-old avian chickens were randomly divided into five groups, and fed on diets as follows: corn-bean control diet (Se 0.2mg/kg) and high Se diets (Se 1mg/kg, high Se group I ; Se 5mg/kg, high Se group II; Se 10mg/kg, high Se group III; Se 15mg/kg, high Se group IV) for 6 weeks. The results were as follow:①The volume, weight and relative weight of thymus, spleen and bursa of Fabricius(BF) were significantly decreased in high Se groups II , III and IV when compared with those of control group. Histopathologically, decreased lymphocytes, enlarged thymic corpuscle and increased nuclear debris in the center of corpuscles were appeared in the thymus. Decreased lymphocytes were observed in spleen and BF of high Se groups II, III and IV. Ultrastructurally, mitochondria injury and increased apoptotic cells with condensed nuclei were appeared in immune organs.②Cell cycle analysis by flow cytometry showed that the G_0G_1 phase was increased, and the S phase, G_2M phase and proliferation index(PI) were decreased in thymus, spleen and BF of high Se II, III and IV. Changes of cell cycle were progressed with the dietary Se level increasing, and consistent with the changes of organs relative weight.③The percentage of Annexin-V positive cells measured by flow cytometry was higher in high Se groups II, III and IV than in control group. TUNEL assay showed that there were increased frequencies of apoptotic cells in high Se groups III and IV. Characteristic of ultrastructural features on apoptotic cells was nuclear membrane breakdown, chromatin condensation with horseshoe-like or crescent shapes. The results were consistent with suppressed growth and decreased lymphocytes of immune organs.④The percentage of CD3+ T cells, CD3+CD4+ T cells and CD3+CD8+ T cells were decreased variably in high Se groups II, III and IV groups when compared with those of control group. The mitogenic reponse of T lymphocytes to ConA and the contents of serum IL-2 were decreased in high Se groups III and IV than in control group. The results suggested that the cellular immune function of chickens was impaired by high Se.⑤The contents of serum IgG were lower in high Se groups III and IV than in control group. The contents of serum IgA and IgM were decreased in high Se group IV. Contents of serum total protein, albumin and globulin were decreased variably in high Se groups III and IV. The results suggested that humoral immune function of chickens was impaired by high Se.⑥The ratio of IC rosette was increased and the ratio of C_3bR rosette decreased in High Se II, III and IV. Immune adherence function of erythrocyte was impaired by high Se.⑦The splenic Se and MDA contents were higher, and splenic GSH-Px, SOD and CAT activities, and total antioxidative ability were lower in high Se groups III and IV than in control group. The structure and function of spleen were impaired. The cellular and humoral function were decreased.
Experiment 2 Mechanisms of the immune functional impairment caused by low Se in chickens
120 one-day-old avian chickens were randomly divided into two groups. Corn, yellow bean and wheat, which were bought from Mianling count were used to formulated low Se diet (Se 0.0342 mg/kg). Sodium selenite was mixed into the low Se diet to produce 0.2 mg/kg Se diet (control group). Broilers were fed the diets for 6 weeks. The results were as follow:①The weight and relative weight of thymus, spleen and BF were significantly decreased in low Se group when compared with those of control group. Anatomically, the volume of immune organs were smaller in low Se group than in control group. Histopathologically, lymphocytes in thymus, spleen and BF were decreased. At 42 days of age, atrophy BF showed infolded epithelium, shrinked follicle with reticuloendothelial cells proliferation. Ultrastructurally, mitochondria injury and increased apoptotic cells with condensed nuclei were observed.②Cell cycle analysis by flow cytometry showed that the G_0G_1 phase was increased, and the G_2M phase and proliferation index(PI) were decreased in thymus, spleen and BF of low Se group. Decreased proliferative function induced growth retardation of immune organs.③Increased apoptotic cells in thymus, spleen and BF were observed by the methods of flow cytometry, TUNEL stain and electron microscope. The results were associated with hypocellular of immune organs.④The percentage of CD3+ T cells and CD3+CD8+ T cells were decreased in low Se group. The mitogenic reponse of T lymphocytes to ConA and the content of serum IL-2 were lower in low Se group than in control group. The results showed that the cellular immune function of chickens was impaired by low Se.⑤The content of serum IgG, IgA, IgM, total protein, albumin and globulin were decreased iIn low Se group. The humoral immune function of chickens were decreased.⑥The ratio of C_3bR rosette was decreased, and the ratio of IC rosette had no obvious change in low Se group. Primary lesion of erythrocyte's adherence function was induced by low Se.⑦The contents of splenic Se, total antioxidative ability, GSH-Px and CAT activities were decreased, and MDA contents in spleen increased in low Se group. The structure and function of spleen were impaired.
The aforementioned results showed that the proliferative function of immune organs and activities of antioxidase in spleen were decreased, and the percentage of apoptotic cells of imunne organs and splenic MDA contents were increased by low Se (0.0342 mg/kg) and high Se (more than 5 mg/kg), which induced growth retardation of immune organs and decrease of cellular immune, humoral immune and erythrocyte's adherence function. The mechanisms of immune injury were closely related to the decreased cellular proliferative function, increased cellular apoptosis, decreased antioxidative ability and accumulated peroxidative products in the immune organs. In the present study, the effect of dietary Se on the structure and function of immune system was systematically researched for the first time. The results provided theoretical basis for determining reasonable supplemental dose of Se and preventing Se-deficiency and selenosis in chickens.
引文
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