硒对山羊妊娠调节的细胞与分子免疫学机制
摘要
硒对动物的免疫与繁殖机能有着重要的影响，探讨硒对妊娠子宫局部细胞因子的调节作用，对阐明硒影响动物繁殖机能的机制，进一步研究其作用和应用具有重要意义。为了从细胞与分子免疫学角度探讨硒对生殖机能调节的机制，进行了以下研究。
试验一：64只成年母兔随机分为3个试验组和1个对照组。试验组分别肌注0.3、0.4和0.5mg·kg-1硒的亚硒酸钠溶液，对照组肌注生理盐水(1.0mL·kg-1)，每周1次，连续8周，复制家兔亚急性硒中毒模型。试验前和给硒2、4、6、8周静脉采血，分析血硒含量。给硒8周后，让公、母兔自然交配，统计发情率、怀孕率及胎仔数。结果表明，试验组血硒浓度显著高于同期对照组(P＜0.05或P＜0.01)，测定血硒浓度可用于硒中毒的临床诊断。硒中毒可引起家兔发情率、怀孕率和胎仔数下降。初步证明，试验所用的硒制剂及其剂量可引起家兔繁殖机能障碍。
试验二：为了研究硒在体外对山羊外周血单个核细胞(peripheral blood mononuclear cell，PBMC)活化的影响，无菌分离未服硒山羊PBMC，分别加入4、8、12和16μg·mL-1的亚硒酸钠进行体外培养，检测硒对PBMC的活化作用及其对分泌IL-2的影响。结果表明，8～12μg·mL-1的亚硒酸钠可显著促进山羊PBMC的活化(P＜0.05或P＜(0.01)且呈剂量依赖关系。4μg·mL-1或16μg·mL-1的亚硒酸钠只能引起PBMC的微弱活化，但与对照组之间差异不显著(P＞0.05)。山羊PBMC体外培养时，加入8～12μg·mL-1的亚硒酸钠可显著促进其IL-2的分泌(P＜0.01)，4μg·mL-1及16μg·mL-1的亚硒酸钠虽能诱导PBMC分泌IL-2，但差异不显著(P＞0.05)。提示在一定范围内，亚硒酸钠能促进山羊PBMC分泌IL-2。其作用呈剂量依赖关系。
试验三：为了分析口服硒对山羊PBMC活化作用的影响，12只母山羊被随机分为A、B、C和D四组。试验A组、试验B和试验C组分别以0.3、0.5和0.7mg·kg-1剂量的硒灌服亚硒酸钠溶液，对照(D)组以1mL·kg-1的蒸馏水灌服，每周1次，连续3周。试验前及每次给硒后24h采血，测定血硒浓度。试验前及每次给硒后一周，无菌采集山羊静脉血抗凝，分离PBMC，体外培养时，用植物血凝素(phytohemagglutinin,PHA-P)刺激，进行活化试验。结果显示：山羊内服0.7mg·kg-1硒2周后；内服0.5和0.7mg·kg-1硒3周后，其PBMC对PHA刺激的反应性显著提高(P＜0.05)。提示山羊内服—定量的硒，可显著提高其免疫细胞对抗原刺激的反应性。
试验四：为了研究硒在体外对妊娠山羊子宫内膜淋巴细胞(endometrial lymphocyte，EML)的活化作用，分离妊娠山羊EML后，分别加入4、8、12和16μg·mL-1的亚硒酸钠进行体外培养，检测硒对EML的活化作用及其对分泌细胞因子IL-2、IL-4、TGFβ1，及TGFβ2的影响。结果显示，8～16μg·mL-1的亚硒酸钠可显著地促进山羊EML的体外活化(P＜0.01)，其活化作用的强弱呈剂量依赖性。EML活化后，对IL-2的分泌呈微弱的促进作用；对IL-4、TGFβ1、TGFβ2，的分泌有显著的促进作用。
汪没乞五：目的是进一步探索硒牙乃在娠早期子宫局部细胞因子的调节作用，妊娠山羊分
为栅A组、B组、Ctg；lt！1对照（D）组。A、B、C理山羊孤以0．3、0．5和0．7呷·哈
剂量的雕Bdjl：I｛jj17qgutIH 溶液，D组灌服IInL·kg‘的蒸馏水，朗1次，濒3周。D组
用邢蔽赚期子宫局部细胞因子IL之、IL4及TGF－日s动态鹏，A、B、C HgHte－－于
观额对n二、R4及TGF－民的调节作用。结果表明：①妊龊期山U于宫角中IL
2、IL4及TGF一 水平无显著差异汐）。②妊娠早期门 d）山羊两侧子宫角
的子宫液中ILZ水平在15～18 d呈上升趋势，19 d时其活性趋于下降。IL4水平一直呈上
升变化。TGF也水平较高，呈不规律变化；TGF也呈现出先升高，后趋于平稳的态势。③
给山羊酮以0．3、0．5和0．7p·kg’剂量的硒灌a哪U顾婪钠蹦一次，渤B3贻，可以
显著陶闲汪娠ibm d子宫液中L－2的水平（尸m历或尸m叫；同时可显著升高IL4的
水平？似伍动PCO贝，胆一定的剂量依赖关系。④给山羊每周以0．5和0．7帖·阶
剂量的硒溶明勋叵西骏钠溶液—次，溯B3贴，可以显著升雕娠Ich18 d子宫液中TGF－
p；衅u ／0 05蔚Pa 01）；每周以 0．7 p·kg’剂量的硒嘲颐匝圃骏钠溶液—次，渤＆3
贻可显著升雕娠Ich17dTGF凡的水平p 05惑0 01），对18d时的TGF＆影响
不显著（a；a砌。
本研究得出以下结论：①首次证明一定量的亚硒酸钠可显著促进山羊EML的体外活
化，并呈一定的剂量依赖关系。EMI活化后，对IL．2的分泌S的促进作用；对IL4、
TGF P；、TGF民的分泌隋显著的促进作用。②首次证明内B勋面刀匣妊娠早期山羊子宫局部
细胞因子thZ 7I＼＝l＝显著陶民脓IL斗、TGF个和TGF－民水钮著升高。今定量的亚
硒酸钠可显著促进山羊PBMC的协【活化，并促进其分泌IL．2。④内BlijBH的硒可显著提
高山羊PBM对PHA的反应性，即细胞免疫功ggte高。⑤中毒量的硒可降低家兔的繁殖
机能。
本研究首次从硒对生殖兔疫的调节作用尤其是对子宫局部细胞因子网络调节作用的角
度，分析阐明了硒对动物繁殖功g髓响的细u。为硒在预防取张浑替夙忙和
提高渊缩额奔仿面的应用提
Selenium(Se) presented significant impact on immunity and reproduction of animals. Research on its regulation to local cytokines secreted in pregnant uterus is of great importance on its regulation mechanism and application to animal reproduction. This study included following five experiments in order to explore the immunological mechanism of its regulation to goat pregnancy at celular and moleacular level.
In Experiment 1 sixty-four adult female rabbits were randomly divided into three experimental groups and a control group. 16 rabbits each. Rabbits in experimental groups were intramuscularly injected with sodium selenite solution respectively at the doses of 0.3. 0.4 and 0.5 mg(Se) ?kg"1 (body weight) and control group with physiological saline (1.0 mL ?kg"'). Each group was treated weekly for eight weeks and then the rabbits in experimental groups acted as subacutely poinsoned model rabbits. Blood samples before treatment and 2. 4, 6 and 8 weeks after treatments were collected from heart or vein and analyzed for selenium content. Female adults in all groups were mated naturally with normal males 8 weeks after treatment. Statistics showed that blood selenium concentration in each experimental group was significantly higher than that of control group (P<0.05 or P<0.01). that the concentration of blood selenium could be clinidally used to diagnose the selenium poisoning, that the toxicity of selenium could cause decrease of oestrus rate, pregnancy rate and the number of fetuses of the births, all proving that the selenious reagent and its doses in this study could result in reproductive barrier in rabbit.
In Experinent 2 the peripheral blood mononuclear cells (PBMC) were aseptically isolated from goats not treated by selenium and cultured in vitro in media containing sodium selenite respectively at the levels of 4, 8. 12 and 16 u g ?mL"'. and then their in
vitro activation and IL-2 levels were tested. The results showed that 8~12 u g ?mL"1 sodium selenite could significantly (P<0.05,orP<0.01) activate goat PBMC in vitro in a dose-dependent manner, that 4 and 16 u g ?mL"' sodium selenite also activated goat PBMC but not significantly(.P>0.05) in comparison to control group (0 u g ?mL"' sodium selenite). The in vitro secretion of IL-2 by goat PBMC was significantly (P<0.01 ) promoted by 8~12 u g -mL"1 sodium selenite but not significantly by 4 and 16 u g ?mL"1 sodium selenite. all indicating that sodium selenite within certain range of doses could dose-dependently stimulate the secretion of IL-2 by goat PBMC.
In Experiment 3 totally 12 female adult goats were randomly arranged in experimental goupes A, B and C and central group D, 3 goats each, in order to analyse the effect of Se orally intaken on the activation of goat PBMC. Goats in group A, B and C orally took sodium selenite solution at doses of 0.3, 0.5 and 0.7 mg(Se)-kg~' respectively and those in control group each orally took 1.0 mL ?kg"1 distilled water once a week for three times. Blood samples were collected from goats before all treatments and 24 h after each treatment and tested for selenium. PBMC were aseptically isolated from goat blood before all treatments and one week after each treatment to conduct in vitro activation, with purified phytohemagglutinin(PHA-P) as activator. The results showed that PBMC from goats 2 weeks after oral administration with 0.7 mg(Se) ?kg"1 or 3 weeks after oral administration with 0.5 and 0.7 mg (Se) ?kg"1 could be significantly (P <0.05~) stimulated by PHA-P, implying that oral administration with a certain amount of Se could sensitize the responseof goat immunocytes to antigen.
Experiment 4 is designed to test the in vitro actvation of endometrial lymphocytes
(EML) by Se. EML were aseptically isolated from pregnant goat, in vitro cultuled in
media respectively containing 4, 8, 12, 16 u g ?mL"1 of sodium selenite and their in vitro
activation and IL-2, IL-4, TGF- P , and TGF- P 2 secretion were assayed. The results
showed that 8?6 u g ?mL"1 sodium selenite significantly activated goat EML in a
试验一：64只成年母兔随机分为3个试验组和1个对照组。试验组分别肌注0.3、0.4和0.5mg·kg-1硒的亚硒酸钠溶液，对照组肌注生理盐水(1.0mL·kg-1)，每周1次，连续8周，复制家兔亚急性硒中毒模型。试验前和给硒2、4、6、8周静脉采血，分析血硒含量。给硒8周后，让公、母兔自然交配，统计发情率、怀孕率及胎仔数。结果表明，试验组血硒浓度显著高于同期对照组(P＜0.05或P＜0.01)，测定血硒浓度可用于硒中毒的临床诊断。硒中毒可引起家兔发情率、怀孕率和胎仔数下降。初步证明，试验所用的硒制剂及其剂量可引起家兔繁殖机能障碍。
试验二：为了研究硒在体外对山羊外周血单个核细胞(peripheral blood mononuclear cell，PBMC)活化的影响，无菌分离未服硒山羊PBMC，分别加入4、8、12和16μg·mL-1的亚硒酸钠进行体外培养，检测硒对PBMC的活化作用及其对分泌IL-2的影响。结果表明，8～12μg·mL-1的亚硒酸钠可显著促进山羊PBMC的活化(P＜0.05或P＜(0.01)且呈剂量依赖关系。4μg·mL-1或16μg·mL-1的亚硒酸钠只能引起PBMC的微弱活化，但与对照组之间差异不显著(P＞0.05)。山羊PBMC体外培养时，加入8～12μg·mL-1的亚硒酸钠可显著促进其IL-2的分泌(P＜0.01)，4μg·mL-1及16μg·mL-1的亚硒酸钠虽能诱导PBMC分泌IL-2，但差异不显著(P＞0.05)。提示在一定范围内，亚硒酸钠能促进山羊PBMC分泌IL-2。其作用呈剂量依赖关系。
试验三：为了分析口服硒对山羊PBMC活化作用的影响，12只母山羊被随机分为A、B、C和D四组。试验A组、试验B和试验C组分别以0.3、0.5和0.7mg·kg-1剂量的硒灌服亚硒酸钠溶液，对照(D)组以1mL·kg-1的蒸馏水灌服，每周1次，连续3周。试验前及每次给硒后24h采血，测定血硒浓度。试验前及每次给硒后一周，无菌采集山羊静脉血抗凝，分离PBMC，体外培养时，用植物血凝素(phytohemagglutinin,PHA-P)刺激，进行活化试验。结果显示：山羊内服0.7mg·kg-1硒2周后；内服0.5和0.7mg·kg-1硒3周后，其PBMC对PHA刺激的反应性显著提高(P＜0.05)。提示山羊内服—定量的硒，可显著提高其免疫细胞对抗原刺激的反应性。
试验四：为了研究硒在体外对妊娠山羊子宫内膜淋巴细胞(endometrial lymphocyte，EML)的活化作用，分离妊娠山羊EML后，分别加入4、8、12和16μg·mL-1的亚硒酸钠进行体外培养，检测硒对EML的活化作用及其对分泌细胞因子IL-2、IL-4、TGFβ1，及TGFβ2的影响。结果显示，8～16μg·mL-1的亚硒酸钠可显著地促进山羊EML的体外活化(P＜0.01)，其活化作用的强弱呈剂量依赖性。EML活化后，对IL-2的分泌呈微弱的促进作用；对IL-4、TGFβ1、TGFβ2，的分泌有显著的促进作用。
汪没乞五：目的是进一步探索硒牙乃在娠早期子宫局部细胞因子的调节作用，妊娠山羊分
为栅A组、B组、Ctg；lt！1对照（D）组。A、B、C理山羊孤以0．3、0．5和0．7呷·哈
剂量的雕Bdjl：I｛jj17qgutIH 溶液，D组灌服IInL·kg‘的蒸馏水，朗1次，濒3周。D组
用邢蔽赚期子宫局部细胞因子IL之、IL4及TGF－日s动态鹏，A、B、C HgHte－－于
观额对n二、R4及TGF－民的调节作用。结果表明：①妊龊期山U于宫角中IL
2、IL4及TGF一 水平无显著差异汐）。②妊娠早期门 d）山羊两侧子宫角
的子宫液中ILZ水平在15～18 d呈上升趋势，19 d时其活性趋于下降。IL4水平一直呈上
升变化。TGF也水平较高，呈不规律变化；TGF也呈现出先升高，后趋于平稳的态势。③
给山羊酮以0．3、0．5和0．7p·kg’剂量的硒灌a哪U顾婪钠蹦一次，渤B3贻，可以
显著陶闲汪娠ibm d子宫液中L－2的水平（尸m历或尸m叫；同时可显著升高IL4的
水平？似伍动PCO贝，胆一定的剂量依赖关系。④给山羊每周以0．5和0．7帖·阶
剂量的硒溶明勋叵西骏钠溶液—次，溯B3贴，可以显著升雕娠Ich18 d子宫液中TGF－
p；衅u ／0 05蔚Pa 01）；每周以 0．7 p·kg’剂量的硒嘲颐匝圃骏钠溶液—次，渤＆3
贻可显著升雕娠Ich17dTGF凡的水平p 05惑0 01），对18d时的TGF＆影响
不显著（a；a砌。
本研究得出以下结论：①首次证明一定量的亚硒酸钠可显著促进山羊EML的体外活
化，并呈一定的剂量依赖关系。EMI活化后，对IL．2的分泌S的促进作用；对IL4、
TGF P；、TGF民的分泌隋显著的促进作用。②首次证明内B勋面刀匣妊娠早期山羊子宫局部
细胞因子thZ 7I＼＝l＝显著陶民脓IL斗、TGF个和TGF－民水钮著升高。今定量的亚
硒酸钠可显著促进山羊PBMC的协【活化，并促进其分泌IL．2。④内BlijBH的硒可显著提
高山羊PBM对PHA的反应性，即细胞免疫功ggte高。⑤中毒量的硒可降低家兔的繁殖
机能。
本研究首次从硒对生殖兔疫的调节作用尤其是对子宫局部细胞因子网络调节作用的角
度，分析阐明了硒对动物繁殖功g髓响的细u。为硒在预防取张浑替夙忙和
提高渊缩额奔仿面的应用提
Selenium(Se) presented significant impact on immunity and reproduction of animals. Research on its regulation to local cytokines secreted in pregnant uterus is of great importance on its regulation mechanism and application to animal reproduction. This study included following five experiments in order to explore the immunological mechanism of its regulation to goat pregnancy at celular and moleacular level.
In Experiment 1 sixty-four adult female rabbits were randomly divided into three experimental groups and a control group. 16 rabbits each. Rabbits in experimental groups were intramuscularly injected with sodium selenite solution respectively at the doses of 0.3. 0.4 and 0.5 mg(Se) ?kg"1 (body weight) and control group with physiological saline (1.0 mL ?kg"'). Each group was treated weekly for eight weeks and then the rabbits in experimental groups acted as subacutely poinsoned model rabbits. Blood samples before treatment and 2. 4, 6 and 8 weeks after treatments were collected from heart or vein and analyzed for selenium content. Female adults in all groups were mated naturally with normal males 8 weeks after treatment. Statistics showed that blood selenium concentration in each experimental group was significantly higher than that of control group (P<0.05 or P<0.01). that the concentration of blood selenium could be clinidally used to diagnose the selenium poisoning, that the toxicity of selenium could cause decrease of oestrus rate, pregnancy rate and the number of fetuses of the births, all proving that the selenious reagent and its doses in this study could result in reproductive barrier in rabbit.
In Experinent 2 the peripheral blood mononuclear cells (PBMC) were aseptically isolated from goats not treated by selenium and cultured in vitro in media containing sodium selenite respectively at the levels of 4, 8. 12 and 16 u g ?mL"'. and then their in
vitro activation and IL-2 levels were tested. The results showed that 8~12 u g ?mL"1 sodium selenite could significantly (P<0.05,orP<0.01) activate goat PBMC in vitro in a dose-dependent manner, that 4 and 16 u g ?mL"' sodium selenite also activated goat PBMC but not significantly(.P>0.05) in comparison to control group (0 u g ?mL"' sodium selenite). The in vitro secretion of IL-2 by goat PBMC was significantly (P<0.01 ) promoted by 8~12 u g -mL"1 sodium selenite but not significantly by 4 and 16 u g ?mL"1 sodium selenite. all indicating that sodium selenite within certain range of doses could dose-dependently stimulate the secretion of IL-2 by goat PBMC.
In Experiment 3 totally 12 female adult goats were randomly arranged in experimental goupes A, B and C and central group D, 3 goats each, in order to analyse the effect of Se orally intaken on the activation of goat PBMC. Goats in group A, B and C orally took sodium selenite solution at doses of 0.3, 0.5 and 0.7 mg(Se)-kg~' respectively and those in control group each orally took 1.0 mL ?kg"1 distilled water once a week for three times. Blood samples were collected from goats before all treatments and 24 h after each treatment and tested for selenium. PBMC were aseptically isolated from goat blood before all treatments and one week after each treatment to conduct in vitro activation, with purified phytohemagglutinin(PHA-P) as activator. The results showed that PBMC from goats 2 weeks after oral administration with 0.7 mg(Se) ?kg"1 or 3 weeks after oral administration with 0.5 and 0.7 mg (Se) ?kg"1 could be significantly (P <0.05~) stimulated by PHA-P, implying that oral administration with a certain amount of Se could sensitize the responseof goat immunocytes to antigen.
Experiment 4 is designed to test the in vitro actvation of endometrial lymphocytes
(EML) by Se. EML were aseptically isolated from pregnant goat, in vitro cultuled in
media respectively containing 4, 8, 12, 16 u g ?mL"1 of sodium selenite and their in vitro
activation and IL-2, IL-4, TGF- P , and TGF- P 2 secretion were assayed. The results
showed that 8?6 u g ?mL"1 sodium selenite significantly activated goat EML in a
引文
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2. Yang FuYu, Wo H. H. Role of selenium in stabilization human erythrocyte membrance skeleton [J].Biochem. Inter. 1987; 16(2):475.
3．徐辉碧．生物微量元素—硒[M]．武汉：华中工学院出版社，1983。
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5．杨克强．硒缺乏对繁殖机能的影响[J]．甘肃畜牧兽医，1995，25(2)：27～28．
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