蒙古马运动相关候选基因的分子鉴定
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摘要
中国是世界上养马历史最悠久的国家之一，但由于机械化的普及和其它马产业的严重滞后，马匹数量和质量发生了严重的下降。近年来，赛马和休闲骑乘在当今世界范围内已然成为一种新兴的产业，国家对于马业重新重视，出台了相关扶持的政策，多地开始新建赛马场和城市乘马俱乐部，并从国外引进多匹良种马。国外的速度赛马多为纯血马，购买和饲养十分昂贵。因此，发展我国的现代马业，必须挖掘我国本土具有优秀运动性能的马品种，通过对其运动性能的充分了解，并自主培育我国赛马，以发展我国赛马种业。
蒙古马是世界上最古老的马品种之一，具有耐力性能卓越、抗严寒、耐粗食和抗病力强等优良特性。并早在13世纪已用于赛马。伴随着分子生物学手段的进步，国外对于纯血马在运动研究的持续开展，对我国特有的地方品种蒙古马进行运动方面的深入详细研究，已迫在眉睫。
本研究以蒙古马为研究对象，研究了与肌肉发育和能量代谢有关基因MSTN、PDK4、CKM、COX411、COX412、PPARGC1A、CAV-1、FABPpm及LDHA在臀中肌表达情况；采用PCR-SSCP及测序的技术对表达量显著差异基因，进行多态性扫描；采用双向电泳和质谱技术鉴定了对不同运动时期的蒙古马肌肉组织差异表达蛋白质。得到的研究结果如下：
1.研究与肌肉发育和能量代谢有关基因MSTN、PDK4、CKM、COX411、COX412、PPARGC1A、CAV-1、FABPpm及LDHA等在蒙古马臀中肌表达模式。
（1）研究蒙古马静止期T0、运动刚刚结束T1和运动后的恢复期T2臀中肌中候选基因表达情况。对上述候选基因的相对表达量进行了测定。结果显示：PDK4和PPARGC1A基因，它们在耐力运动后mRNA的表达量都发生了上调，在运动后的恢复期mRNA的表达量水平甚至还发生了显著的上调（P<0.05），且二者具有显著的相关性；蒙古马CAV-1基因的mRNA的表达量在运动后的恢复期发生了显著的上调（P<0.05），但运动刚刚结束相对运动前的静止期略有减少。
（2）对训练骑乘过的蒙古马与未曾训练的蒙古马静止状态下臀中肌中候选基因mRNA的表达量检测后发现，训练对蒙古马所选候选基因的mRNA的表达量均有影响，其中PDK4、MSTN及COX412三个基因mRNA的表达量发生了显著的增加，特别是COX412基因发生了极显著的增加（P<0.01）。
（3）采用Western blotting的方法对PDK4、PGC-1和CAV-1蛋白在蒙古马不同运动时期表达量进行测定，结果显示：PDK4、CAV-1和PGC-1三个蛋白在耐力运动后较运动前的静止期均有增加，且PDK4在运动后恢复期蛋白表达量较运动前有了显著的增加（P<0.05），而另外两个蛋白增加并未到达显著水平。
2.利用差异候选基因法，确定对在蒙古马在耐力运动前后候选基因mRNA的表达量显著差异的CAV-1，PDK4和PPARGC1A三个基因为蒙古马耐力运动的候选基因，采用SSCP-PCR和直接测序的方法进行基因外显子区多态研究，结果表明三个基因在蒙古马中高度保守，仅在PPARGC1A第五外显子区发现单核苷酸多态。从群体遗传学的角度分析统计了4个品种马PPARGC1A基因第五外显子PCR-SSCP位点的基因型频率和等位基因频率，发现等位基因A比B占优势，AA基因型频率比AB和BB占优势。
3.利用双向电泳的方法对蒙古马不同运动时期骨骼肌中的蛋白质组学进行研究，结果发现不同运动时期共有蛋白差异点44个，其中运动刚刚结束与运动前静止期有31个差异蛋白质点，运动后恢复期与运动前静止期比较有30个差异蛋白点。经MALDI一TOF/MS质谱分析，成功鉴定44个不同运动时期差异表达蛋白点，分值高于78阈值（P<0.05）的不同蛋白质有28个。对这些蛋白进行GO注释及WEGO分析，结果显示：若按生物过程分，包含有16类，其中代谢过程占35.7%，包括ACADVL、HSPB1、HSP70、LARS、TNNT3和Rab-40c等10个蛋白。
China is one of the oldest countries to have horse raising, but the quantity andquality of horse have declined seriously because of the popularity of mechanization andserious lagging of other horse industries. In recent years, horse racing and leisure horseriding have became emerging industries around the world, so horse industry have beenaroused attention again by a great of states, and some relevant supporting policies havebeen introduced. Many places started to build racing courses and city saddle horse clubsand imported many thoroughbred horses from abroad. Most of the foreign racing horsesare thoroughbred horses which are too expensive to buy and breed. It’s necessary to dignational horse breeds which have good motion performance by fully understanding theirlocomotion, and to breed our national racing horse and develop Chinese racing horseindustry.
Mongolian horse is one of the oldest horse breeds in the world, and has powerfulendurance, strong resistance to chilliness, rough feeding and disease. And it has beenused for racing from the13th century to nowadays. Along with the progress of molecularbiology methods and continuous studying of thoroughbred horses on their locomotion byforeign countries, it’s extremely urgent to study the area of sports of Mongolian horsewhich is unique local breed in our country.
In this study, we studied the expression of muscle development and energymetabolism related genes including MSTN, PDK4, CKM, COX411, COX412,PPARGC1A, CAV-1, FABPpm and LDHA gene in gluteus medius of Mongolian horse;PCR-SSCP and sequencing technology were used to carry out polymorphism scanningfor genes which showed significantly different expression; Two-dimensionalelectrophoresis and mass spectrometry were used to identify differential expressiveproteins in muscular tissues of Mongolian horses from different periods of movement.We got the following results:
1. Studying the expressive pattern of muscle development related genes includingMSTN, PDK4, CKM, COX411, COX412, PPARGC1A, CAV-1, FABPpm and LDHA genein gluteus medius of Mongolian horses.
(1) Studying the expression of candidate genes in gluteus medius at stationary phaseT0, movement immediately postexercise phase T1and recovery phase T2from movementand measuring the relative expression of these candidate genes. The results showed that: the mRNA expressions of PDK4and PPARGC1A gene were up regulate at phase T1andsignificantly up regulate (P<0.05) at phase T2, the both have significant relativity; themRNA expression of CAV-1gene was significantly up regulate (P<0.05) at phase T2,while expression at phase T1was lower than phase T0.
(2) After measuring the mRNA expressions of the candidate genes in gluteus mediusof trained Mongolian horses and untrained Mongolian horses at stationary condition, wefound training had effects on mRNA expressions of all candidate genes, within raletivegenes the mRNA expressions of PDK4, MSTN and COX412gene increased significantly(P<0.05) and COX412gene was significant higher (P<0.01).
(3) Western blotting was used to measure the protein expressions of PDK4, PGC-1and CAV-1gene of Mongolian horses at different periods of movement, the resultsshowed protein expressions of PDK4, CAV-1and PGC-1gene were higher at phase T1and T2than phase T0, and protein expression of PDK4gene was significant higher(P<0.05) at phase T2than phase T0, while the other two genes didn’t reach significantlevel.
2. Choosing CAV-1, PDK4and PPARGC1A gene as candidate genes for enduranceexercise of Mongolian horses because the mRNA expressions of these genes weresignificantly different before and after endurance exercise. SSCP-PCR was used to studythe polymorphisms in exons of the genes, the results showed the three genes were highlyconserved in Mongolian horses and just found single nucleotide polymorphisms in thefifth exon of PPARGC1A gene. And then we detected PCR-SSCP of PPARGC1A gene infour horse populations, and found allele A was the dominant allele.
3. Two-dimensional electrophoresis was used to study proteomics of skeletal muscleof Mongolian horses at different periods of movement, we found44different proteinpoints at different movement phases, within there were31different protein pointsbetween phase T1and phase T0,30different protein points between phase T2and phaseT0. We successfully identified44different protein points in different movement stagesand got28different proteins by MALDI-TOF/MS mass spectrometry. These proteins areGO noted and WEGO analysied. The results showed that there are16classes accodingbiology processes, consist metabolic processes (35.7%), mainly include10proteins, suchas ACADVL、HSPB1、HSP70、LARS、TNNT3、Rab-40c, and so forth.
蒙古马是世界上最古老的马品种之一，具有耐力性能卓越、抗严寒、耐粗食和抗病力强等优良特性。并早在13世纪已用于赛马。伴随着分子生物学手段的进步，国外对于纯血马在运动研究的持续开展，对我国特有的地方品种蒙古马进行运动方面的深入详细研究，已迫在眉睫。
本研究以蒙古马为研究对象，研究了与肌肉发育和能量代谢有关基因MSTN、PDK4、CKM、COX411、COX412、PPARGC1A、CAV-1、FABPpm及LDHA在臀中肌表达情况；采用PCR-SSCP及测序的技术对表达量显著差异基因，进行多态性扫描；采用双向电泳和质谱技术鉴定了对不同运动时期的蒙古马肌肉组织差异表达蛋白质。得到的研究结果如下：
1.研究与肌肉发育和能量代谢有关基因MSTN、PDK4、CKM、COX411、COX412、PPARGC1A、CAV-1、FABPpm及LDHA等在蒙古马臀中肌表达模式。
（1）研究蒙古马静止期T0、运动刚刚结束T1和运动后的恢复期T2臀中肌中候选基因表达情况。对上述候选基因的相对表达量进行了测定。结果显示：PDK4和PPARGC1A基因，它们在耐力运动后mRNA的表达量都发生了上调，在运动后的恢复期mRNA的表达量水平甚至还发生了显著的上调（P<0.05），且二者具有显著的相关性；蒙古马CAV-1基因的mRNA的表达量在运动后的恢复期发生了显著的上调（P<0.05），但运动刚刚结束相对运动前的静止期略有减少。
（2）对训练骑乘过的蒙古马与未曾训练的蒙古马静止状态下臀中肌中候选基因mRNA的表达量检测后发现，训练对蒙古马所选候选基因的mRNA的表达量均有影响，其中PDK4、MSTN及COX412三个基因mRNA的表达量发生了显著的增加，特别是COX412基因发生了极显著的增加（P<0.01）。
（3）采用Western blotting的方法对PDK4、PGC-1和CAV-1蛋白在蒙古马不同运动时期表达量进行测定，结果显示：PDK4、CAV-1和PGC-1三个蛋白在耐力运动后较运动前的静止期均有增加，且PDK4在运动后恢复期蛋白表达量较运动前有了显著的增加（P<0.05），而另外两个蛋白增加并未到达显著水平。
2.利用差异候选基因法，确定对在蒙古马在耐力运动前后候选基因mRNA的表达量显著差异的CAV-1，PDK4和PPARGC1A三个基因为蒙古马耐力运动的候选基因，采用SSCP-PCR和直接测序的方法进行基因外显子区多态研究，结果表明三个基因在蒙古马中高度保守，仅在PPARGC1A第五外显子区发现单核苷酸多态。从群体遗传学的角度分析统计了4个品种马PPARGC1A基因第五外显子PCR-SSCP位点的基因型频率和等位基因频率，发现等位基因A比B占优势，AA基因型频率比AB和BB占优势。
3.利用双向电泳的方法对蒙古马不同运动时期骨骼肌中的蛋白质组学进行研究，结果发现不同运动时期共有蛋白差异点44个，其中运动刚刚结束与运动前静止期有31个差异蛋白质点，运动后恢复期与运动前静止期比较有30个差异蛋白点。经MALDI一TOF/MS质谱分析，成功鉴定44个不同运动时期差异表达蛋白点，分值高于78阈值（P<0.05）的不同蛋白质有28个。对这些蛋白进行GO注释及WEGO分析，结果显示：若按生物过程分，包含有16类，其中代谢过程占35.7%，包括ACADVL、HSPB1、HSP70、LARS、TNNT3和Rab-40c等10个蛋白。
China is one of the oldest countries to have horse raising, but the quantity andquality of horse have declined seriously because of the popularity of mechanization andserious lagging of other horse industries. In recent years, horse racing and leisure horseriding have became emerging industries around the world, so horse industry have beenaroused attention again by a great of states, and some relevant supporting policies havebeen introduced. Many places started to build racing courses and city saddle horse clubsand imported many thoroughbred horses from abroad. Most of the foreign racing horsesare thoroughbred horses which are too expensive to buy and breed. It’s necessary to dignational horse breeds which have good motion performance by fully understanding theirlocomotion, and to breed our national racing horse and develop Chinese racing horseindustry.
Mongolian horse is one of the oldest horse breeds in the world, and has powerfulendurance, strong resistance to chilliness, rough feeding and disease. And it has beenused for racing from the13th century to nowadays. Along with the progress of molecularbiology methods and continuous studying of thoroughbred horses on their locomotion byforeign countries, it’s extremely urgent to study the area of sports of Mongolian horsewhich is unique local breed in our country.
In this study, we studied the expression of muscle development and energymetabolism related genes including MSTN, PDK4, CKM, COX411, COX412,PPARGC1A, CAV-1, FABPpm and LDHA gene in gluteus medius of Mongolian horse;PCR-SSCP and sequencing technology were used to carry out polymorphism scanningfor genes which showed significantly different expression; Two-dimensionalelectrophoresis and mass spectrometry were used to identify differential expressiveproteins in muscular tissues of Mongolian horses from different periods of movement.We got the following results:
1. Studying the expressive pattern of muscle development related genes includingMSTN, PDK4, CKM, COX411, COX412, PPARGC1A, CAV-1, FABPpm and LDHA genein gluteus medius of Mongolian horses.
(1) Studying the expression of candidate genes in gluteus medius at stationary phaseT0, movement immediately postexercise phase T1and recovery phase T2from movementand measuring the relative expression of these candidate genes. The results showed that: the mRNA expressions of PDK4and PPARGC1A gene were up regulate at phase T1andsignificantly up regulate (P<0.05) at phase T2, the both have significant relativity; themRNA expression of CAV-1gene was significantly up regulate (P<0.05) at phase T2,while expression at phase T1was lower than phase T0.
(2) After measuring the mRNA expressions of the candidate genes in gluteus mediusof trained Mongolian horses and untrained Mongolian horses at stationary condition, wefound training had effects on mRNA expressions of all candidate genes, within raletivegenes the mRNA expressions of PDK4, MSTN and COX412gene increased significantly(P<0.05) and COX412gene was significant higher (P<0.01).
(3) Western blotting was used to measure the protein expressions of PDK4, PGC-1and CAV-1gene of Mongolian horses at different periods of movement, the resultsshowed protein expressions of PDK4, CAV-1and PGC-1gene were higher at phase T1and T2than phase T0, and protein expression of PDK4gene was significant higher(P<0.05) at phase T2than phase T0, while the other two genes didn’t reach significantlevel.
2. Choosing CAV-1, PDK4and PPARGC1A gene as candidate genes for enduranceexercise of Mongolian horses because the mRNA expressions of these genes weresignificantly different before and after endurance exercise. SSCP-PCR was used to studythe polymorphisms in exons of the genes, the results showed the three genes were highlyconserved in Mongolian horses and just found single nucleotide polymorphisms in thefifth exon of PPARGC1A gene. And then we detected PCR-SSCP of PPARGC1A gene infour horse populations, and found allele A was the dominant allele.
3. Two-dimensional electrophoresis was used to study proteomics of skeletal muscleof Mongolian horses at different periods of movement, we found44different proteinpoints at different movement phases, within there were31different protein pointsbetween phase T1and phase T0,30different protein points between phase T2and phaseT0. We successfully identified44different protein points in different movement stagesand got28different proteins by MALDI-TOF/MS mass spectrometry. These proteins areGO noted and WEGO analysied. The results showed that there are16classes accodingbiology processes, consist metabolic processes (35.7%), mainly include10proteins, suchas ACADVL、HSPB1、HSP70、LARS、TNNT3、Rab-40c, and so forth.
引文
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