不同硒源对鸡原代肝细胞硒酶活性及mRNA表达的影响
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摘要
硒是人和动物必需的微量元素,具有提高机体免疫力、抗癌、抗自由基、延缓衰老、拮抗有毒元素、防治某些地方性流行病等多种功能。许多国家都将日粮补硒作为预防动物缺硒的常规措施。在动物体内,硒存在的主要形式是硒蛋白。多数硒蛋白又是具有重要作用的酶,称之为硒酶。对硒酶活性和基因表达的研究已经获得了许多重要成果,但仍有不少尚未澄清的问题。本文在建立了鸡原代肝细胞无血清培养模型和荧光定量PCR检测方法的基础上,研究了不同浓度的硒卡拉胶(Se-Car)、亚硒酸钠(Na-Se)和硒蛋氨酸(Se-Met)对鸡肝细胞谷胱甘肽过氧化物酶(GPx1)活性及磷脂氢谷胱甘肽过氧化物酶(GPx4)和脱碘酶Ⅰ(IDⅠ)mRNA表达的影响,为不同硒源的作用机理提供理论依据。
试验1鸡肝细胞中GPx4和IDⅠmRNA水平荧光定量PCR检测方法的建立根据已知的鸡β-actin、GPx4和IDⅠ基因的mRNA序列,设计并合成3对特异性引物。从鸡肝细胞中抽提总RNA,反转录,合成cDNA,再以cDNA为模板,分别进行3个目的基因片段的PCR(polymerase chain reaction,PCR)扩增。胶回收PCR产物,测定浓度并10倍梯度稀释作为建立荧光定量PCR的标准模板,同时,对引物浓度和退火温度进行优化。结果显示,3对引物各扩增出一条目的带,大小和预期相符合.反应的最佳引物浓度是10 pmol·μL~(-1),最佳退火温度为58℃。扩增β-actin、GPx4和IDⅠ基因的3对引物的扩增效率分别为93.5%、91.4%和98.4%,且PCR产物的熔解曲线显示单一峰。
试验2鸡原代肝细胞分离及无血清培养方法的建立采用胶原酶灌注法,对30-40日龄的白来航鸡进行肝细胞分离,用台盼蓝拒染法计算肝细胞总数和实时存活率。肝细胞以1×10~6个·ml~(-1)接种于6板,每孔2.5ml,无血清L-15培养基,37℃普通恒温箱培养。观察细胞形态,测定不同培养时间细胞培养基上清液中的乳酸脱氢酶(LDH)活力。结果显示,每个鸡肝脏分离获得的细胞总量为(4.1±0.2)×10~8个,肝细胞实时存活率为92%±1%;LDH活力在培养后4--96 h之间呈下降趋势,细胞可维持良好的生长状态达6天以上。
试验3不同硒源对鸡原代肝细胞GPx1活性的影响肝细胞培养24h后,将三种硒源分别以0(对照),0.5,1,1.5,2,3,4,5μmol·L~(-1)的浓度(以Se计)加入到培养液中,继续培养24h。测定培养基上清液中LDH活力,细胞内谷胱甘肽(GSH)含量和GPx1活力。结果显示:LDH活力随硒浓度增加而增加,但是当Se-Met在1-3μmol·L~(-1)时,LDH活力低于对照组(P＜0.05);GSH含量随硒浓度增加先降后升,Se-Car浓度在1.5-2.0μmol·L~(-1)(P＜0.01)、Se-Met浓度在1.0-2.0μmol·L~(-1)(P＜0.01)、Na-Se浓度在1.5μmol·L~(-1)(P＜0.05)时,GSH的浓度与对照组相比显著降低,Se-Car浓度在4.0-5.0μmol·L~(-1)(P＜0.01)、Se-Met浓度在4.0-5.0μmol·L~(-1)(P＜0.05)、Na-Se浓度在3.0-5.0μmol·L~(-1)(P＜0.01)时,GSH浓度与对照组相比显著升高;GPx1活力随硒浓度增加先升后降,当Se-Car和Se-Met浓度为2μmol·L~(-1)、Na-Se浓度为1.5μmol·L~(-1)时,各组GPx1活力达到最大(P＜0.001),当Se-Car和Se-Met浓度在2-5μmol·L~(-1)区间、Na-Se浓度在1.5-5μmol·L~(-1)区间时,GPx1活力呈下降趋势。结论:一定浓度范围内,Se-Met比Se-Car和Na-Se具有更大的安全范围和更好的抗氧化能力。
试验4不同硒源对鸡原代肝细胞IDⅠ及GPx4 mRNA表达的影响试验处理同试验3,然后测定IDⅠ和GPx4 mRNA表达水平。结果显示:三种硒源添加组IDⅠmRNA表达水平均显著高于对照(P＜0.05),随硒浓度增加IDⅠmRNA表达水平下降,下降幅度Se-Met＜Na-Se＜Se-Car。三种硒源添加组GPx4 mRNA表达水平均极显著低于对照组(P＜0.001),随硒浓度增加GPx4 mRNA表达水平下降,下降幅度Se-Met＜Se-Car＜Na-Se。结论:一定浓度范围内,三种硒源均可显著提高IDⅠmRNA表达水平,Se-Met在较宽的浓度范围内可使IDⅠmRNA在高水平维持,Na-Se和Se-Car次之;三种硒源均可使GPx4 mRNA表达水平显著下降,下降幅度Se-Met更小。
Selenium is an essential trace element for animals and human.Its functions include improving immunity,preventing cancer,resisting free radicals,postponing aging, detoxification from some heavy metals,and preventing some local epidemic etc.In many countries,Supplementation of selenium in animal diet is a routine step for preventing Se shortage on animal husbandry.Selenoproteins,most of which are also called selenoenzymes that possess important functions in organism,are the main selenium forms in animals.Many achievements have been obtained in the acitivity and gene expression study of selenoenzymes,however,there still are many unclear problems to solve.In this paper,the serum-free culture for primary chicken hepatocytes and the real-time PCR method for measuring mRNA levels of GPx4 and IDⅠwere developed and then the effect of different concentrations of Se-car,Se-Met and Na-Se on the activity of GPx1 and the mRNA expressions of GPx4 and IDⅠin primary cultured chicken hepatocytes was investigated.
Experiment 1.Development of method for determination of GPx4 and IDⅠmRNA levels in chicken hepatocytes.According to the published mRNA sequences of gene ofβ-actin,GPx4 and IDⅠ,three pairs of specific primers were designed by biological software and synthesized.Total RNA was extracted from chicken liver and the cDNA was synthesised using Oligo dT primer.Using the synthesised cDNA as templates,three target fragments were amplified by polymerase chain reaction(PCR),respectively.The PCR products were electrophoresed on 2%agarose gels and extracted.The concentrations of purified PCR products were measured and diluted by 10-folds.Real-time PCRs were performed based on the 10-fold dilution series of standard DNA templates.In addition,the contents and annealing temperatures for the three pairs of specific primers were optimized, respectively.The results showed that one target band,which was consistent with the fact, was amplified for each specific primer pair.The optimal contents and annealing temperatures were 10μmol/L and 58℃,respectively.Amplification efficiencies of the primer pairs for gene ofβ-actin,GPx4 and IDⅠwere 93.5%,91.4%and 98.4%, respectively.The melting curve analysis showed only one peak for each PCR product.
Experiment 2.Development of isolation and serum-free culture methods for primary chicken hepatocytes.Chicken hepatocytes were obtained from white Leghorn chicken(30～40-day old) by collagenase perfusion.Total cell count and survival rate of freshly isolated hepatocytes were obtained by applying trypan blue exclusion.Hepatocytes were seeded into a 6-well microplate at a density of 2.5×10~6 cells/2.5ml/well and cultured at 37℃with free atmosphere exchange.Morphological assessment of hepatocyte monolayers and determination of LDH activity in the culture supernatant were performed. The results showed that the cell yield was(4.1±0.2)×10~8 hepatocytes per liver.The survival rate of freshly isolated hepatocytes was 92%±1%.Cultured in this condition,the LDH activity decreased gradually from 4h to 96h after seeding and the chicken hepatocytes could maintain good state up to 6 days.
Experiment 3.Effect of different concentrations of Se-Car,Na-Se,and Se-Met on GPx1 activity in primary cultured chicken hepatocytes.Chicken hepatocytes were cultured for 24 h,then the old media were removed and the cells were washed three times and cultured in fresh media supplemented with 0(control),0.5,1,1.5,2,3,4,or 5μmol·L~(-1) Se as Se-Car,Na-Se,or Se-Met for another 24 h.Activities of LDH in culture media,GPx1 activities and concentrations of GSH in hepatocyte cytosol were measured.The results showed that the LDH activities rised with the increased doses of Se,however,the activities of LDH in the culture media supplemented with Se-Met at doses of 1-3μmol/L were significantly lower than those of controls(P＜0.05).When compared to controls, concentrations of GSH decreased significantly at 1.5-2μmol·L~(-1) Se-Car(P＜0.01),1.5μmol·L~(-1) Na-Se(P＜0.05),and 1-2μmol·L~(-1) Se-Met(P＜0.01),while GSH concentrations increased significantly at 4-5μmol·L~(-1) Se-Car(P＜0.01),3-5μmol·L~(-1) Na-Se(P＜0.01), and 4-5μmol·L~(-1) Se-Met(P＜0.05),respectively.Activities of GPx1 reached the maximums at 2μmol·L~(-1) Se-Car(2.18 folds vs.control,P＜0.001),1.5μmol·L~(-1) Na-Se (2.16 folds vs.controls,P＜0.001),and 2μmol·L~(-1) Se-Met(2.48 folds vs.control,P＜0.001).Trends of decrease in GPx1 activities were observed at 2-5μmol·L~(-1) Se-Car,1.5-5μmol·L~(-1) Na-Se,and 2-5μmol·L~(-1) Se-Met,in a dose-dependent manner.Our results indicated that Se-Met was better than Se-Car and Na-Se in terms of security range and antioxidant ability.
Experiment 4.Effect of different concentrations of Se-Car,Se-Met and Na-Se on the mRNA expression of IDⅠand GPx4 in primary cultured chicken hepatocytes.The treatment of chicken hepatocytes was the same as that in experiment 3.The mRNA levels were measured by real-time PCR.For Se-Car and Na-Se groups,the highest levels of mRNA for IDⅠwere observed at 0.5-1.5μmol·L~(-1) and 0.5μmol·L~(-1),respectively,then they decresed dose-dependently.For Se-Met groups,the IDⅠmRNA levels increased at concentrations of 0.5-1.0μmol·L~(-1),kept high and steady at concentrations of 0.5-1.0μmol·L~(-1) and then decreased slightly.For each concentration of the three selenium sources, GPx4 mRNA levels declined very significantly(P＜0.001 vs.control).With the increase of selenium,GPx4 mRNA levels declined with no difference in Se-Met groups,whereas decreased significantly in the rest two.This suggested that compared with the rest two, Se-Met could keep a high level of mRNA for IDⅠin wider concentration range.All of the three could decrease the expression of GPx4 mRNA significantly with less decrease in Se-Met groups than in the rest two.
试验1鸡肝细胞中GPx4和IDⅠmRNA水平荧光定量PCR检测方法的建立根据已知的鸡β-actin、GPx4和IDⅠ基因的mRNA序列,设计并合成3对特异性引物。从鸡肝细胞中抽提总RNA,反转录,合成cDNA,再以cDNA为模板,分别进行3个目的基因片段的PCR(polymerase chain reaction,PCR)扩增。胶回收PCR产物,测定浓度并10倍梯度稀释作为建立荧光定量PCR的标准模板,同时,对引物浓度和退火温度进行优化。结果显示,3对引物各扩增出一条目的带,大小和预期相符合.反应的最佳引物浓度是10 pmol·μL~(-1),最佳退火温度为58℃。扩增β-actin、GPx4和IDⅠ基因的3对引物的扩增效率分别为93.5%、91.4%和98.4%,且PCR产物的熔解曲线显示单一峰。
试验2鸡原代肝细胞分离及无血清培养方法的建立采用胶原酶灌注法,对30-40日龄的白来航鸡进行肝细胞分离,用台盼蓝拒染法计算肝细胞总数和实时存活率。肝细胞以1×10~6个·ml~(-1)接种于6板,每孔2.5ml,无血清L-15培养基,37℃普通恒温箱培养。观察细胞形态,测定不同培养时间细胞培养基上清液中的乳酸脱氢酶(LDH)活力。结果显示,每个鸡肝脏分离获得的细胞总量为(4.1±0.2)×10~8个,肝细胞实时存活率为92%±1%;LDH活力在培养后4--96 h之间呈下降趋势,细胞可维持良好的生长状态达6天以上。
试验3不同硒源对鸡原代肝细胞GPx1活性的影响肝细胞培养24h后,将三种硒源分别以0(对照),0.5,1,1.5,2,3,4,5μmol·L~(-1)的浓度(以Se计)加入到培养液中,继续培养24h。测定培养基上清液中LDH活力,细胞内谷胱甘肽(GSH)含量和GPx1活力。结果显示:LDH活力随硒浓度增加而增加,但是当Se-Met在1-3μmol·L~(-1)时,LDH活力低于对照组(P＜0.05);GSH含量随硒浓度增加先降后升,Se-Car浓度在1.5-2.0μmol·L~(-1)(P＜0.01)、Se-Met浓度在1.0-2.0μmol·L~(-1)(P＜0.01)、Na-Se浓度在1.5μmol·L~(-1)(P＜0.05)时,GSH的浓度与对照组相比显著降低,Se-Car浓度在4.0-5.0μmol·L~(-1)(P＜0.01)、Se-Met浓度在4.0-5.0μmol·L~(-1)(P＜0.05)、Na-Se浓度在3.0-5.0μmol·L~(-1)(P＜0.01)时,GSH浓度与对照组相比显著升高;GPx1活力随硒浓度增加先升后降,当Se-Car和Se-Met浓度为2μmol·L~(-1)、Na-Se浓度为1.5μmol·L~(-1)时,各组GPx1活力达到最大(P＜0.001),当Se-Car和Se-Met浓度在2-5μmol·L~(-1)区间、Na-Se浓度在1.5-5μmol·L~(-1)区间时,GPx1活力呈下降趋势。结论:一定浓度范围内,Se-Met比Se-Car和Na-Se具有更大的安全范围和更好的抗氧化能力。
试验4不同硒源对鸡原代肝细胞IDⅠ及GPx4 mRNA表达的影响试验处理同试验3,然后测定IDⅠ和GPx4 mRNA表达水平。结果显示:三种硒源添加组IDⅠmRNA表达水平均显著高于对照(P＜0.05),随硒浓度增加IDⅠmRNA表达水平下降,下降幅度Se-Met＜Na-Se＜Se-Car。三种硒源添加组GPx4 mRNA表达水平均极显著低于对照组(P＜0.001),随硒浓度增加GPx4 mRNA表达水平下降,下降幅度Se-Met＜Se-Car＜Na-Se。结论:一定浓度范围内,三种硒源均可显著提高IDⅠmRNA表达水平,Se-Met在较宽的浓度范围内可使IDⅠmRNA在高水平维持,Na-Se和Se-Car次之;三种硒源均可使GPx4 mRNA表达水平显著下降,下降幅度Se-Met更小。
Selenium is an essential trace element for animals and human.Its functions include improving immunity,preventing cancer,resisting free radicals,postponing aging, detoxification from some heavy metals,and preventing some local epidemic etc.In many countries,Supplementation of selenium in animal diet is a routine step for preventing Se shortage on animal husbandry.Selenoproteins,most of which are also called selenoenzymes that possess important functions in organism,are the main selenium forms in animals.Many achievements have been obtained in the acitivity and gene expression study of selenoenzymes,however,there still are many unclear problems to solve.In this paper,the serum-free culture for primary chicken hepatocytes and the real-time PCR method for measuring mRNA levels of GPx4 and IDⅠwere developed and then the effect of different concentrations of Se-car,Se-Met and Na-Se on the activity of GPx1 and the mRNA expressions of GPx4 and IDⅠin primary cultured chicken hepatocytes was investigated.
Experiment 1.Development of method for determination of GPx4 and IDⅠmRNA levels in chicken hepatocytes.According to the published mRNA sequences of gene ofβ-actin,GPx4 and IDⅠ,three pairs of specific primers were designed by biological software and synthesized.Total RNA was extracted from chicken liver and the cDNA was synthesised using Oligo dT primer.Using the synthesised cDNA as templates,three target fragments were amplified by polymerase chain reaction(PCR),respectively.The PCR products were electrophoresed on 2%agarose gels and extracted.The concentrations of purified PCR products were measured and diluted by 10-folds.Real-time PCRs were performed based on the 10-fold dilution series of standard DNA templates.In addition,the contents and annealing temperatures for the three pairs of specific primers were optimized, respectively.The results showed that one target band,which was consistent with the fact, was amplified for each specific primer pair.The optimal contents and annealing temperatures were 10μmol/L and 58℃,respectively.Amplification efficiencies of the primer pairs for gene ofβ-actin,GPx4 and IDⅠwere 93.5%,91.4%and 98.4%, respectively.The melting curve analysis showed only one peak for each PCR product.
Experiment 2.Development of isolation and serum-free culture methods for primary chicken hepatocytes.Chicken hepatocytes were obtained from white Leghorn chicken(30～40-day old) by collagenase perfusion.Total cell count and survival rate of freshly isolated hepatocytes were obtained by applying trypan blue exclusion.Hepatocytes were seeded into a 6-well microplate at a density of 2.5×10~6 cells/2.5ml/well and cultured at 37℃with free atmosphere exchange.Morphological assessment of hepatocyte monolayers and determination of LDH activity in the culture supernatant were performed. The results showed that the cell yield was(4.1±0.2)×10~8 hepatocytes per liver.The survival rate of freshly isolated hepatocytes was 92%±1%.Cultured in this condition,the LDH activity decreased gradually from 4h to 96h after seeding and the chicken hepatocytes could maintain good state up to 6 days.
Experiment 3.Effect of different concentrations of Se-Car,Na-Se,and Se-Met on GPx1 activity in primary cultured chicken hepatocytes.Chicken hepatocytes were cultured for 24 h,then the old media were removed and the cells were washed three times and cultured in fresh media supplemented with 0(control),0.5,1,1.5,2,3,4,or 5μmol·L~(-1) Se as Se-Car,Na-Se,or Se-Met for another 24 h.Activities of LDH in culture media,GPx1 activities and concentrations of GSH in hepatocyte cytosol were measured.The results showed that the LDH activities rised with the increased doses of Se,however,the activities of LDH in the culture media supplemented with Se-Met at doses of 1-3μmol/L were significantly lower than those of controls(P＜0.05).When compared to controls, concentrations of GSH decreased significantly at 1.5-2μmol·L~(-1) Se-Car(P＜0.01),1.5μmol·L~(-1) Na-Se(P＜0.05),and 1-2μmol·L~(-1) Se-Met(P＜0.01),while GSH concentrations increased significantly at 4-5μmol·L~(-1) Se-Car(P＜0.01),3-5μmol·L~(-1) Na-Se(P＜0.01), and 4-5μmol·L~(-1) Se-Met(P＜0.05),respectively.Activities of GPx1 reached the maximums at 2μmol·L~(-1) Se-Car(2.18 folds vs.control,P＜0.001),1.5μmol·L~(-1) Na-Se (2.16 folds vs.controls,P＜0.001),and 2μmol·L~(-1) Se-Met(2.48 folds vs.control,P＜0.001).Trends of decrease in GPx1 activities were observed at 2-5μmol·L~(-1) Se-Car,1.5-5μmol·L~(-1) Na-Se,and 2-5μmol·L~(-1) Se-Met,in a dose-dependent manner.Our results indicated that Se-Met was better than Se-Car and Na-Se in terms of security range and antioxidant ability.
Experiment 4.Effect of different concentrations of Se-Car,Se-Met and Na-Se on the mRNA expression of IDⅠand GPx4 in primary cultured chicken hepatocytes.The treatment of chicken hepatocytes was the same as that in experiment 3.The mRNA levels were measured by real-time PCR.For Se-Car and Na-Se groups,the highest levels of mRNA for IDⅠwere observed at 0.5-1.5μmol·L~(-1) and 0.5μmol·L~(-1),respectively,then they decresed dose-dependently.For Se-Met groups,the IDⅠmRNA levels increased at concentrations of 0.5-1.0μmol·L~(-1),kept high and steady at concentrations of 0.5-1.0μmol·L~(-1) and then decreased slightly.For each concentration of the three selenium sources, GPx4 mRNA levels declined very significantly(P＜0.001 vs.control).With the increase of selenium,GPx4 mRNA levels declined with no difference in Se-Met groups,whereas decreased significantly in the rest two.This suggested that compared with the rest two, Se-Met could keep a high level of mRNA for IDⅠin wider concentration range.All of the three could decrease the expression of GPx4 mRNA significantly with less decrease in Se-Met groups than in the rest two.
引文
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