丙酮酸肌酸对肉鸡脂肪代谢和蛋白质代谢的影响及其机制研究
摘要
丙酮酸肌酸(creatine pyruvate, CrPyr)是一种新的功能营养素,其对丙酮酸和肌酸的生物学功能进行了有效的结合和互补。丙酮酸肌酸的主要功能是抑制肌肉中蛋白质的流失、增加肌肉重量,同时促进脂肪代谢、减少体脂百分数。目前,对于CrPyr功能的研究主要集中在运动生理学方面。其在畜禽生产方面,特别是对家禽脂肪代谢和蛋白质代谢影响的研究鲜有报道。本研究以雄性AA肉鸡为研究对象,探讨日粮添加丙酮酸肌酸对肉鸡脂肪代谢及蛋白质代谢的影响；并以比较蛋白质组学为主要研究手段,构建CrPyr处理的肉鸡肝脏和肌肉组织蛋白表达谱,分析表达差异并揭示各相关蛋白质/酶之间的作用规律和相互关系,揭示丙酮酸肌酸调节肉鸡脂肪代谢及蛋白质代谢生物化学机制。
1日粮添加不同剂量的丙酮酸肌酸对肉鸡脂肪代谢和蛋白质代谢的影响
为探讨丙酮酸肌酸(CrPyr)对肉鸡脂肪代谢和蛋白质代谢的影响,400羽1日龄雄性AA肉鸡随机分为4组,每组4个重复,每个重复25羽。按常规饲养方式饲喂至22日龄,分别于基础日粮中添加0%、1%、5%和10% CrPyr。饲喂至42日龄,各组分别从每个重复中随机抽取3只肉鸡宰杀,采集血样、肝脏、腹脂等冻存,待测。试剂盒法检测肝脏甘油三酯(TG)、血清TG、总胆固醇(TC)、高密度脂蛋白胆固醇(HDL-C)、低密度脂蛋白胆固醇(LDL-C)、非酯化脂肪酸(NEFA)和尿酸(UA)的含量及肌酸激酶(CK)的活性；放射免疫分析法检测血清中胰高血糖素(Glucagon)、胰岛素(Insulin)和瘦素(Leptin)的含量；荧光实时定量PCR法检测肝脏组织中脂肪代谢关键因子乙酰辅酶A羧化酶(ACC)、脂肪酸合酶(FAS)、肉碱棕榈酰转移酶-I(CPT-I)、过氧化物酶体增殖物激活受体a (PPAR-a)及肌肉中胰岛素样生长因子-1 (IGF-1)、肌肉中生肌调节因子(Myogenin)、肌肉生长抑制素(Myostatin)基因的mRNA表达水平。试验结果显示,10% CrPyr显著降低肉鸡体重(BW)和日增重(ADG)。5%和10% CrPyr均可以显著降低肉鸡的腹脂率(AFR)、血清及肝脏中甘油三酯(TG)含量。相反,5%和10% CrPyr显著升高相对胸肌重量、血清中NEFA、HDL-C、Glucagon. Insulin和Leptin含量及CK活性。基因表达分析结果显示,5%和10% CrPyr可促进肝脏PPAR-a、CPT-I和肌肉IGF-1的mRNA表达,降低肌肉Myostatin的mRNA表达。结果提示,添加5%的CrPyr在不影响生长性能的情况下,能够降低AA肉鸡脂肪的沉积,同时增强体蛋白的沉积。
2丙酮酸、丙酮酸肌酸和肌酸对肉鸡脂肪代谢及蛋白质代谢的影响
为比较丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡脂肪代谢和蛋白质代谢的影响,400羽1日龄雄性AA肉鸡随机分为4组,每组4个重复,每个重复25羽。按常规饲养方式饲喂至22日龄,对照组饲喂基础日粮,试验组分别于基础日粮中添加5% CrPyr、2% Pyr和3%Cr。饲喂至42日龄时,各组分别从每个重复中随机随机抽取3只肉鸡宰杀,采集血样、肝脏、腹脂等冻存,待测。检测指标同试验一试验结果发现,5% CrPyr对肉鸡的体重及饲料转化率并无影响；CrPyr和Pyr均显著降低肝脏TG和血清TC; CrPyr显著升高相对腿肌重量、血清中NEFA和HDL-C的含量以及肝脏组织中CPT-I和PPAR-α基因的mRNA表达水平；CrPyr和Cr均显著降低血清中UA的含量和肌肉组织中Myostatin基因的mRNA表达水平,显著升高肌肉组织中IGF-1基因的mRNA表达水平。结果提示,CrPyr在降低肉鸡的脂肪代谢沉积和增强蛋白质沉积方面均强于Pyr和Cr,体现了其双功能营养素的作用效应。
3丙酮酸、丙酮酸肌酸和肌酸对肉鸡肝脏线粒体蛋白/酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,研究丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肝脏线粒体脂肪代谢相关蛋白/酶表达谱的影响,拟从蛋白水平探讨Pyr, CrPyr和Cr调节肉鸡脂肪代谢和蛋白质代谢的机理。试验设计同第二章。试验方法采用多步差速离心和密度梯度离心联合法分离纯化肝脏线粒体。采用测定溶酶体标志酶(半乳糖苷酶),质膜标志酶(碱性磷酸二酯酶),线粒体标志酶(细胞色素C氧化酶)和胞浆标志酶(乳酸脱氢酶)的活性对线粒体纯度评价；以琥珀酸脱氢酶活性估算线粒体的富集度；以及用专一的荧光探针(MitoTracker RED)观察线粒体形态。结果表明,利用差速离心和等密度梯度离心的联合可以有效地分离纯化线粒体。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组得到13个差异蛋白,Pyr处理组得到17个差异蛋白,Cr处理组得到11个差异蛋白。试验结果表明,CrPyr通过下调脂肪分化相关蛋白(ADRP)的表达减少脂肪酸的聚积,抑制ATP合酶的表达从而减少脂肪细胞的生成,同时通过下调胆固醇酯转移蛋白(CETP)的表达提高HDL水平和降低TG水平,减少脂肪和胆固醇的沉积；CrPyr和Cr上调蛋白质合成过程中起关键作用的翻译起始因子2B (eIF2B)因子的蛋白水平,促进蛋白质的合成,且CrPyr作用效果强于Cr; Pyr和Cr上调脂酰CoA合成酶(ACS)的蛋白表达,促进脂肪酸的β-氧化分解,减少脂肪沉积；Pyr上调参与蛋白质终止后加工、转运的甲硫氨酸胺基肽酶、鸟氨酸转移酶和热休克蛋白60的表达。与Pyr和Cr比较,CrPyr在脂肪分解和蛋白质合成上发挥了更为重要的作用,且作用效果均强于Pyr和Cr。
4丙酮酸、丙酮酸肌酸和肌酸对肉鸡肝脏胞浆蛋白/酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,探讨丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肝脏胞浆蛋白/酶表达的影响,拟从蛋白水平探讨Pyr, CrPyr和Cr调节肉鸡脂肪代谢和蛋白质代谢的机理。试验设计同第二章。试验方法采用差速离心法分离肝脏胞浆蛋白。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组得到16个差异蛋白,Pyr处理组得到19个差异蛋白,Cr处理组得到12个差异蛋白。试验结果表明,CrPyr和Pyr通过上调肉鸡肝脏胞浆中脂肪酸结合蛋白(FABP)、下调胆固醇酯转移蛋白(CETP)的表达,加快脂肪酸的氧化分解和TG的分解,减少脂肪的沉积,且CrPyr作用效果比Pyr显著；CrPyr和Cr通过上调载脂蛋白A-IV (ApoA-IV)的表达,加快TG的分解,减少脂肪的堆积,且CrPyr作用效果比Cr显著；CyPry还通过上调肉鸡肝脏胞浆中钙网织蛋白(CRT)和真核翻译起始因子3a (eIF3a)的水平,促进蛋白质的合成,而Pyr和Cr对肉鸡肝脏的蛋白质代谢没有明显的影响。因此,同Pyr和Cr相比较,CrPyr在肉鸡肝脏脂肪代谢和蛋白质代谢方面发挥了更积极有效的作用。
5丙酮酸、丙酮酸肌酸和肌酸对肉鸡肌肉蛋白／酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,探讨丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肌肉蛋白／酶表达谱的影响,拟从蛋白质水平探讨Pyr, CrPyr和Cr调节肉鸡肌肉蛋白质代谢的机理。试验设计同第二章。试验方法采用组织匀浆法分离肌肉蛋白。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组鉴定出11个差异蛋白,Pyr处理组鉴定出11个差异蛋白,Cr处理组鉴定出得到10个差异蛋白。试验结果表明,CrPyr通过增加肌肉中肌酸激酶(MCK)的活性,升高磷酸肌酸的含量；同时降低磷酸化酶b激酶的活性,减少糖元的分解,使肌肉能量储备增强,进而影响机体的能量代谢。CrPyr上调真核翻译起始因子2(eIF2)的表达水平,提示其可以促进肌肉蛋白质的合成,最终表现为肌肉组织中肌球蛋白轻链和肌动蛋白的表达的增加。Pyr上调糖酵解过程中的关键酶丙酮酸激酶(PK)和烯醇化酶的表达,同时下调参与肌细胞内参与氧运输和贮存的肌红蛋白的表达,提示Pyr可以加快缺氧条件下的糖元酵解,对机体能量的恢复有一定的帮助,而对肌肉的蛋白质代谢没有影响。Cr对肌球蛋白和肌动蛋白的表达有所影响,同时可以提高肌酸激酶的活性,但是表达量低于CrPyr处理组。综合比较显示,CrPyr在肌肉能量代谢和蛋白质代谢上发挥了积极的作用,且作用效果明显高于其单独的有效成分Cr和Pyr。
Creatine pyruvate (CrPyr) is a new multifunctional nutrient, it makes a very good combination and complement with the function of pyruvate and creatine. The mainly biological functions of CrPyr are inhibite the loss of muscle protein, increase muscle mass and promote lipid metabolism, reduce body fat percentage and delay fatigue. Currently, the research for CrPyr focused on exercise physiology, its lipid and protein metabolism in broiler chickens has been reported rarely. In this study, a study was carried out to examine the effects of exogenous added CrPyr on lipid and protein metabolism in male AA broilers; Through the changes in protein profiles of liver and muscle by using comparative proteomics, the purpose of this is to reveal the relationship between the proteins and analyse the mechanism of CrPyr effects on lipid and protein metabolism in broiler chickens.
1 Effect of CrPyr with different doses on lipid and protein metabolism in broiler chickens
To assess the effects of creatine pyruvate (CrPyr) on lipid and protein metabolism in broiler chickens, a total of 400 one day-old male birds (Arbor Acres) were randomly allocated to four groups, with each group replicated four times and each replicate involving 25 birds. The broilers were provided with a commercial diet supplemented with CrPyr at 0%,1%,5% or 10% of the diet, respectively, for a period of 3 wks ad libitum (from 22 to 42 d). The content of hepatic triglycerides (TG), blood triglycerides (TG), total cholesterol (TC), high density lipoprotein-cholesterol (HDL-C), low density lipoprotein-cholesterol (LDL-C), non-esterified fatty acid (NEFA), uric acid (UA), creatine kinase (CK), glucagon (Glu), insulin (Ins), leptin (Lep), triiodothyronine (T3), thyroxin (T4), free triiodothyronine (FT3) and free thyroxin (FT4) were determined by using commercial kits. The genes of acetyl CoA carboxylase (ACC), fatty acid synthetase (FAS), peroxisome proliferators activated receptor a (PPAR-a), carnitine palmitoyl transferase-I (CPT-I) in liver and muscle insulin-like growth factor 1 (IGF-1), Myostatin, Myogenin in muscle for real-time PCR assays. Four broiler chickens per replicate from each treatment were chosen randomly and slaughtered to collect specimens of blood, liver, abdominal fat at 42 day-old. In the present study, body weight (BW) and average daily gain (ADG) of broilers decreased in 10% CrPyr group.5% or 10% CrPyr of diets decreased the abdominal fat rate (AFR, abdominal fat/live weight), serum or hepatic TG concentrations of the broilers. In contrast,5% and 10% CrPyr caused a marked increase in the relative pectoral muscle weights, serum NEFA, HDL-C, Glu, Ins or Lep contents and CK activity,. The expression of hepatic PPAR-αand CPT-I, muscle IGF-1 were significantly elevated in the 5% and 10% groups, and the Myostatin mRNA expression was reduced. It was found that supplementation with 5% CrPyr enhances lipolysis and protein synthesis, while not adversely affects growth performance in broiler chickens.
2 Comparison of Pyr, CrPyr and Cr on lipid and protein metabolism in broiler chickens
The effects of Pyruvate (Pyr), Creatine Pyruvate (CrPyr) and Creatine (Cr) on lipid and protein metabolism were compared in broiler chickens. Four hundred one day-old male birds (Arbor Acres) were allocated to four groups, each of which included four replicates (25 birds per replicate). Treatments consisted of unsupplemented basal diet (0%), basal diet containing 2% Pyr, basal diet containing 5% CrPyr and basal diet containing3% Cr, respectively, for a period of 3 wks ad libitum (from 22 to 42 d). Three broiler chickens per replicate from each treatment were chosen randomly and slaughtered to collect specimens of blood, liver, abdominal fat at 42 day-old. The detected index were same as text one. The results showed that CrPyr and Pyr significantly decreased the hepatic TG and serum TC concentration. CrPyr markedly increased the relative leg muscle weights, the serum NEFA and HDL-C concentrations, while the expression of CPT-I and PPAR-a mRNA in the liver were both decidedly enhanced in the CrPyr group. The serum UA content was and the Myostatin mRNA level was reduced in the CrPyr and Cr groups. Muscle IGF-1 expression was enhanced in both the CrPyr and Cr groups. In addition, CrPyr did not alter body weight or the feed conversion ratio. These results indicate that, compared with Pyr and Cr alone, CrPyr has a bifunctional role in broiler chickens, in that it influences both lipid and protein metabolism.
3 Effects of Pyr, CrPyr and Cr on the expressions of liver mitochondrial protein/ enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of liver mitochondrial lipid and protein metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapte. Mitochondrial purification process using the two-step differential centrifugation and density gradient method. The activities of galactosidase (lysome), alkaline phosphodiesterase (plasma membrane), cytochrome c oxidase (mitochondria), and lactate dehydrogenase were determined to assess the purity of mitochondrion; the activities of fumaric reductase was determined to estimate the enrichment of mitochondrion; Otherwise the morphous of mitochondrion was observed by using the MitoTracker RED. The results showed that mitochondrion was separated and purificated efficiently by using this methord in present study. Proteins were isolated by differential centrifugation and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 13 different proteins, Pyr treatment group received 17 different proteins, Cr treatment group received 11 different proteins. The results showed that, CrPyr reduced the accumulation of fatty acids by down-regulating the expression of ADRP, inhibited the expression of ATP synthase to reduce the formation of fat cells, while reduced the expression of CETP to increase HDL levels and decrease TG levels, resulting in the less fat and cholesterol deposition; CrPyr and Cr increased the expression of eIF2B, which played a key role in protein synthesis, thus promoted protein synthesis, and the effect of CrPyr was more significantly than Cr; Pyr and Cr raised the expression of ACS to promote theβ-oxidation of fatty acids and reduced fat deposition; Pyr also raised the levels of methionine aminopeptidase, ornithine aminotransferase and 60kDa heat shock protein, which effected on protein processing and transport after the termination of protein synthesis. Compared with the Pyr and Cr, CrPyr plays a more important role in lipolysis and protein synthesis, and the effect was stronger than Pyr and Cr.
4 Effects of Pyr, CrPyr and Cr on the expressions of liver cytosolic protein/enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of liver cytosolic lipid and protein metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapter. Proteins were isolated by differential centrifugation and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 16 different proteins, Pyr treatment group received 19 different proteins, Cr treatment group received 12 different proteins. The results showed that, CrPyr and Pyr up-regulated the expression of FABP and CETP in liver cytoplasm to accelerate the decomposition of fatty acids and TG, thus reducing fat deposition, the effects of CrPyr was more significantly than Pyr; CrPyr and Cr raised the expression of ApoA-IV, it could accelerate the decomposition levels of TG, then reduced fat accumulation, the effects of CrPyr was significantly than Cr; Also, CyPry increased the levels of CRT and eIF3a to promote protein synthesis, and Pyr and Cr did not have effects on protein metabolism in chicken liver. Therefore, compared with Pyr and Cr, CrPyr plays a more active and effective role in lipid and protein metabolism in broiler chickens.
5 Effects of Pyr, CrPyr and Cr on the expressions of muscle protein/enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of muscle metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapter. Proteins were isolated by homogenates and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, and then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 11 different proteins, Pyr treatment group received 11 different proteins, Cr treatment group received 10 different proteins. The results showed that, CrPyr increased the activity of MCK, elevated the levels of creatine phosphate; it also reduced the activity of phosphorylase b kinase to decrease the decomposition of glycogen and increase the body's energy reserves, thereby affecting the body's energy metabolism. CrPyr up-regulated the expression of eIF2, this suggested that it could promote muscle protein synthesis, the final performance were the increased expression of muscle myosin light chain and actin. Pyr raised the expression of PK and enolase, which were the key enzyme in glycolysis, while reduced the expression of myoglobin, which played a role in the transport and storage of oxygen in muscle cells, suggesting that Pyr could help the energy recovery of the body through glycolysis under hypoxia accelerated; Pyr did not have effects on muscle protein metabolism. Cr changed the expression of myosin and actin, while increased the activity of creatine kinase, but the expression was lower than CrPyr treatment group. Comprehensive comparison shows that, CrPyr plays an active role in muscle energy metabolism and protein metabolism and the effect was significantly higher than the individual active ingredients Cr and Pyr.
1日粮添加不同剂量的丙酮酸肌酸对肉鸡脂肪代谢和蛋白质代谢的影响
为探讨丙酮酸肌酸(CrPyr)对肉鸡脂肪代谢和蛋白质代谢的影响,400羽1日龄雄性AA肉鸡随机分为4组,每组4个重复,每个重复25羽。按常规饲养方式饲喂至22日龄,分别于基础日粮中添加0%、1%、5%和10% CrPyr。饲喂至42日龄,各组分别从每个重复中随机抽取3只肉鸡宰杀,采集血样、肝脏、腹脂等冻存,待测。试剂盒法检测肝脏甘油三酯(TG)、血清TG、总胆固醇(TC)、高密度脂蛋白胆固醇(HDL-C)、低密度脂蛋白胆固醇(LDL-C)、非酯化脂肪酸(NEFA)和尿酸(UA)的含量及肌酸激酶(CK)的活性；放射免疫分析法检测血清中胰高血糖素(Glucagon)、胰岛素(Insulin)和瘦素(Leptin)的含量；荧光实时定量PCR法检测肝脏组织中脂肪代谢关键因子乙酰辅酶A羧化酶(ACC)、脂肪酸合酶(FAS)、肉碱棕榈酰转移酶-I(CPT-I)、过氧化物酶体增殖物激活受体a (PPAR-a)及肌肉中胰岛素样生长因子-1 (IGF-1)、肌肉中生肌调节因子(Myogenin)、肌肉生长抑制素(Myostatin)基因的mRNA表达水平。试验结果显示,10% CrPyr显著降低肉鸡体重(BW)和日增重(ADG)。5%和10% CrPyr均可以显著降低肉鸡的腹脂率(AFR)、血清及肝脏中甘油三酯(TG)含量。相反,5%和10% CrPyr显著升高相对胸肌重量、血清中NEFA、HDL-C、Glucagon. Insulin和Leptin含量及CK活性。基因表达分析结果显示,5%和10% CrPyr可促进肝脏PPAR-a、CPT-I和肌肉IGF-1的mRNA表达,降低肌肉Myostatin的mRNA表达。结果提示,添加5%的CrPyr在不影响生长性能的情况下,能够降低AA肉鸡脂肪的沉积,同时增强体蛋白的沉积。
2丙酮酸、丙酮酸肌酸和肌酸对肉鸡脂肪代谢及蛋白质代谢的影响
为比较丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡脂肪代谢和蛋白质代谢的影响,400羽1日龄雄性AA肉鸡随机分为4组,每组4个重复,每个重复25羽。按常规饲养方式饲喂至22日龄,对照组饲喂基础日粮,试验组分别于基础日粮中添加5% CrPyr、2% Pyr和3%Cr。饲喂至42日龄时,各组分别从每个重复中随机随机抽取3只肉鸡宰杀,采集血样、肝脏、腹脂等冻存,待测。检测指标同试验一试验结果发现,5% CrPyr对肉鸡的体重及饲料转化率并无影响；CrPyr和Pyr均显著降低肝脏TG和血清TC; CrPyr显著升高相对腿肌重量、血清中NEFA和HDL-C的含量以及肝脏组织中CPT-I和PPAR-α基因的mRNA表达水平；CrPyr和Cr均显著降低血清中UA的含量和肌肉组织中Myostatin基因的mRNA表达水平,显著升高肌肉组织中IGF-1基因的mRNA表达水平。结果提示,CrPyr在降低肉鸡的脂肪代谢沉积和增强蛋白质沉积方面均强于Pyr和Cr,体现了其双功能营养素的作用效应。
3丙酮酸、丙酮酸肌酸和肌酸对肉鸡肝脏线粒体蛋白/酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,研究丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肝脏线粒体脂肪代谢相关蛋白/酶表达谱的影响,拟从蛋白水平探讨Pyr, CrPyr和Cr调节肉鸡脂肪代谢和蛋白质代谢的机理。试验设计同第二章。试验方法采用多步差速离心和密度梯度离心联合法分离纯化肝脏线粒体。采用测定溶酶体标志酶(半乳糖苷酶),质膜标志酶(碱性磷酸二酯酶),线粒体标志酶(细胞色素C氧化酶)和胞浆标志酶(乳酸脱氢酶)的活性对线粒体纯度评价；以琥珀酸脱氢酶活性估算线粒体的富集度；以及用专一的荧光探针(MitoTracker RED)观察线粒体形态。结果表明,利用差速离心和等密度梯度离心的联合可以有效地分离纯化线粒体。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组得到13个差异蛋白,Pyr处理组得到17个差异蛋白,Cr处理组得到11个差异蛋白。试验结果表明,CrPyr通过下调脂肪分化相关蛋白(ADRP)的表达减少脂肪酸的聚积,抑制ATP合酶的表达从而减少脂肪细胞的生成,同时通过下调胆固醇酯转移蛋白(CETP)的表达提高HDL水平和降低TG水平,减少脂肪和胆固醇的沉积；CrPyr和Cr上调蛋白质合成过程中起关键作用的翻译起始因子2B (eIF2B)因子的蛋白水平,促进蛋白质的合成,且CrPyr作用效果强于Cr; Pyr和Cr上调脂酰CoA合成酶(ACS)的蛋白表达,促进脂肪酸的β-氧化分解,减少脂肪沉积；Pyr上调参与蛋白质终止后加工、转运的甲硫氨酸胺基肽酶、鸟氨酸转移酶和热休克蛋白60的表达。与Pyr和Cr比较,CrPyr在脂肪分解和蛋白质合成上发挥了更为重要的作用,且作用效果均强于Pyr和Cr。
4丙酮酸、丙酮酸肌酸和肌酸对肉鸡肝脏胞浆蛋白/酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,探讨丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肝脏胞浆蛋白/酶表达的影响,拟从蛋白水平探讨Pyr, CrPyr和Cr调节肉鸡脂肪代谢和蛋白质代谢的机理。试验设计同第二章。试验方法采用差速离心法分离肝脏胞浆蛋白。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组得到16个差异蛋白,Pyr处理组得到19个差异蛋白,Cr处理组得到12个差异蛋白。试验结果表明,CrPyr和Pyr通过上调肉鸡肝脏胞浆中脂肪酸结合蛋白(FABP)、下调胆固醇酯转移蛋白(CETP)的表达,加快脂肪酸的氧化分解和TG的分解,减少脂肪的沉积,且CrPyr作用效果比Pyr显著；CrPyr和Cr通过上调载脂蛋白A-IV (ApoA-IV)的表达,加快TG的分解,减少脂肪的堆积,且CrPyr作用效果比Cr显著；CyPry还通过上调肉鸡肝脏胞浆中钙网织蛋白(CRT)和真核翻译起始因子3a (eIF3a)的水平,促进蛋白质的合成,而Pyr和Cr对肉鸡肝脏的蛋白质代谢没有明显的影响。因此,同Pyr和Cr相比较,CrPyr在肉鸡肝脏脂肪代谢和蛋白质代谢方面发挥了更积极有效的作用。
5丙酮酸、丙酮酸肌酸和肌酸对肉鸡肌肉蛋白／酶表达谱的影响
本试验选取雄性AA肉鸡为研究对象,探讨丙酮酸(Pyr)、丙酮酸肌酸(CrPyr)和肌酸(Cr)对肉鸡肌肉蛋白／酶表达谱的影响,拟从蛋白质水平探讨Pyr, CrPyr和Cr调节肉鸡肌肉蛋白质代谢的机理。试验设计同第二章。试验方法采用组织匀浆法分离肌肉蛋白。通过2D-PAGE分离后,经图谱分析获得差异蛋白点,再进行MALDI-TOF-MS质谱鉴定。与对照组相比,CrPyr处理组鉴定出11个差异蛋白,Pyr处理组鉴定出11个差异蛋白,Cr处理组鉴定出得到10个差异蛋白。试验结果表明,CrPyr通过增加肌肉中肌酸激酶(MCK)的活性,升高磷酸肌酸的含量；同时降低磷酸化酶b激酶的活性,减少糖元的分解,使肌肉能量储备增强,进而影响机体的能量代谢。CrPyr上调真核翻译起始因子2(eIF2)的表达水平,提示其可以促进肌肉蛋白质的合成,最终表现为肌肉组织中肌球蛋白轻链和肌动蛋白的表达的增加。Pyr上调糖酵解过程中的关键酶丙酮酸激酶(PK)和烯醇化酶的表达,同时下调参与肌细胞内参与氧运输和贮存的肌红蛋白的表达,提示Pyr可以加快缺氧条件下的糖元酵解,对机体能量的恢复有一定的帮助,而对肌肉的蛋白质代谢没有影响。Cr对肌球蛋白和肌动蛋白的表达有所影响,同时可以提高肌酸激酶的活性,但是表达量低于CrPyr处理组。综合比较显示,CrPyr在肌肉能量代谢和蛋白质代谢上发挥了积极的作用,且作用效果明显高于其单独的有效成分Cr和Pyr。
Creatine pyruvate (CrPyr) is a new multifunctional nutrient, it makes a very good combination and complement with the function of pyruvate and creatine. The mainly biological functions of CrPyr are inhibite the loss of muscle protein, increase muscle mass and promote lipid metabolism, reduce body fat percentage and delay fatigue. Currently, the research for CrPyr focused on exercise physiology, its lipid and protein metabolism in broiler chickens has been reported rarely. In this study, a study was carried out to examine the effects of exogenous added CrPyr on lipid and protein metabolism in male AA broilers; Through the changes in protein profiles of liver and muscle by using comparative proteomics, the purpose of this is to reveal the relationship between the proteins and analyse the mechanism of CrPyr effects on lipid and protein metabolism in broiler chickens.
1 Effect of CrPyr with different doses on lipid and protein metabolism in broiler chickens
To assess the effects of creatine pyruvate (CrPyr) on lipid and protein metabolism in broiler chickens, a total of 400 one day-old male birds (Arbor Acres) were randomly allocated to four groups, with each group replicated four times and each replicate involving 25 birds. The broilers were provided with a commercial diet supplemented with CrPyr at 0%,1%,5% or 10% of the diet, respectively, for a period of 3 wks ad libitum (from 22 to 42 d). The content of hepatic triglycerides (TG), blood triglycerides (TG), total cholesterol (TC), high density lipoprotein-cholesterol (HDL-C), low density lipoprotein-cholesterol (LDL-C), non-esterified fatty acid (NEFA), uric acid (UA), creatine kinase (CK), glucagon (Glu), insulin (Ins), leptin (Lep), triiodothyronine (T3), thyroxin (T4), free triiodothyronine (FT3) and free thyroxin (FT4) were determined by using commercial kits. The genes of acetyl CoA carboxylase (ACC), fatty acid synthetase (FAS), peroxisome proliferators activated receptor a (PPAR-a), carnitine palmitoyl transferase-I (CPT-I) in liver and muscle insulin-like growth factor 1 (IGF-1), Myostatin, Myogenin in muscle for real-time PCR assays. Four broiler chickens per replicate from each treatment were chosen randomly and slaughtered to collect specimens of blood, liver, abdominal fat at 42 day-old. In the present study, body weight (BW) and average daily gain (ADG) of broilers decreased in 10% CrPyr group.5% or 10% CrPyr of diets decreased the abdominal fat rate (AFR, abdominal fat/live weight), serum or hepatic TG concentrations of the broilers. In contrast,5% and 10% CrPyr caused a marked increase in the relative pectoral muscle weights, serum NEFA, HDL-C, Glu, Ins or Lep contents and CK activity,. The expression of hepatic PPAR-αand CPT-I, muscle IGF-1 were significantly elevated in the 5% and 10% groups, and the Myostatin mRNA expression was reduced. It was found that supplementation with 5% CrPyr enhances lipolysis and protein synthesis, while not adversely affects growth performance in broiler chickens.
2 Comparison of Pyr, CrPyr and Cr on lipid and protein metabolism in broiler chickens
The effects of Pyruvate (Pyr), Creatine Pyruvate (CrPyr) and Creatine (Cr) on lipid and protein metabolism were compared in broiler chickens. Four hundred one day-old male birds (Arbor Acres) were allocated to four groups, each of which included four replicates (25 birds per replicate). Treatments consisted of unsupplemented basal diet (0%), basal diet containing 2% Pyr, basal diet containing 5% CrPyr and basal diet containing3% Cr, respectively, for a period of 3 wks ad libitum (from 22 to 42 d). Three broiler chickens per replicate from each treatment were chosen randomly and slaughtered to collect specimens of blood, liver, abdominal fat at 42 day-old. The detected index were same as text one. The results showed that CrPyr and Pyr significantly decreased the hepatic TG and serum TC concentration. CrPyr markedly increased the relative leg muscle weights, the serum NEFA and HDL-C concentrations, while the expression of CPT-I and PPAR-a mRNA in the liver were both decidedly enhanced in the CrPyr group. The serum UA content was and the Myostatin mRNA level was reduced in the CrPyr and Cr groups. Muscle IGF-1 expression was enhanced in both the CrPyr and Cr groups. In addition, CrPyr did not alter body weight or the feed conversion ratio. These results indicate that, compared with Pyr and Cr alone, CrPyr has a bifunctional role in broiler chickens, in that it influences both lipid and protein metabolism.
3 Effects of Pyr, CrPyr and Cr on the expressions of liver mitochondrial protein/ enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of liver mitochondrial lipid and protein metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapte. Mitochondrial purification process using the two-step differential centrifugation and density gradient method. The activities of galactosidase (lysome), alkaline phosphodiesterase (plasma membrane), cytochrome c oxidase (mitochondria), and lactate dehydrogenase were determined to assess the purity of mitochondrion; the activities of fumaric reductase was determined to estimate the enrichment of mitochondrion; Otherwise the morphous of mitochondrion was observed by using the MitoTracker RED. The results showed that mitochondrion was separated and purificated efficiently by using this methord in present study. Proteins were isolated by differential centrifugation and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 13 different proteins, Pyr treatment group received 17 different proteins, Cr treatment group received 11 different proteins. The results showed that, CrPyr reduced the accumulation of fatty acids by down-regulating the expression of ADRP, inhibited the expression of ATP synthase to reduce the formation of fat cells, while reduced the expression of CETP to increase HDL levels and decrease TG levels, resulting in the less fat and cholesterol deposition; CrPyr and Cr increased the expression of eIF2B, which played a key role in protein synthesis, thus promoted protein synthesis, and the effect of CrPyr was more significantly than Cr; Pyr and Cr raised the expression of ACS to promote theβ-oxidation of fatty acids and reduced fat deposition; Pyr also raised the levels of methionine aminopeptidase, ornithine aminotransferase and 60kDa heat shock protein, which effected on protein processing and transport after the termination of protein synthesis. Compared with the Pyr and Cr, CrPyr plays a more important role in lipolysis and protein synthesis, and the effect was stronger than Pyr and Cr.
4 Effects of Pyr, CrPyr and Cr on the expressions of liver cytosolic protein/enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of liver cytosolic lipid and protein metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapter. Proteins were isolated by differential centrifugation and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 16 different proteins, Pyr treatment group received 19 different proteins, Cr treatment group received 12 different proteins. The results showed that, CrPyr and Pyr up-regulated the expression of FABP and CETP in liver cytoplasm to accelerate the decomposition of fatty acids and TG, thus reducing fat deposition, the effects of CrPyr was more significantly than Pyr; CrPyr and Cr raised the expression of ApoA-IV, it could accelerate the decomposition levels of TG, then reduced fat accumulation, the effects of CrPyr was significantly than Cr; Also, CyPry increased the levels of CRT and eIF3a to promote protein synthesis, and Pyr and Cr did not have effects on protein metabolism in chicken liver. Therefore, compared with Pyr and Cr, CrPyr plays a more active and effective role in lipid and protein metabolism in broiler chickens.
5 Effects of Pyr, CrPyr and Cr on the expressions of muscle protein/enzyme in broiler chickens
The male birds (Arbor Acres) were selected to study the effect of Pyr, CrPyr and Cr on the expressions of muscle metabolism-related protein/enzyme in broiler chickens. Experimental design was same to the second chapter. Proteins were isolated by homogenates and were separated by two-dimensional electrophoresis, the differentially displayed spots were obtained by pattern analysis, and then the spots were further identified by MALDI-TOF-MS. Compared with the control group, CrPyr treatment group received 11 different proteins, Pyr treatment group received 11 different proteins, Cr treatment group received 10 different proteins. The results showed that, CrPyr increased the activity of MCK, elevated the levels of creatine phosphate; it also reduced the activity of phosphorylase b kinase to decrease the decomposition of glycogen and increase the body's energy reserves, thereby affecting the body's energy metabolism. CrPyr up-regulated the expression of eIF2, this suggested that it could promote muscle protein synthesis, the final performance were the increased expression of muscle myosin light chain and actin. Pyr raised the expression of PK and enolase, which were the key enzyme in glycolysis, while reduced the expression of myoglobin, which played a role in the transport and storage of oxygen in muscle cells, suggesting that Pyr could help the energy recovery of the body through glycolysis under hypoxia accelerated; Pyr did not have effects on muscle protein metabolism. Cr changed the expression of myosin and actin, while increased the activity of creatine kinase, but the expression was lower than CrPyr treatment group. Comprehensive comparison shows that, CrPyr plays an active role in muscle energy metabolism and protein metabolism and the effect was significantly higher than the individual active ingredients Cr and Pyr.
引文
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