牦牛肉宰后成熟过程中组织蛋白酶L、B、H对肉品质及微观结构的影响
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摘要
肉的可接受性又被称为肉的食用品质，是消费者和肉类生产者共同关注的问题。在肉的食用品质中，嫩度是决定肉品质的重要指标。大量研究证明，对动物宰后胴体进行低温冷却成熟可以提高肉的嫩度。肉类工作者一致认为肉的成熟过程中嫩度的改变是内源性蛋白酶(钙激活酶系、蛋白酶体及组织蛋白酶系)作用的结果。本文以10头青海大通牦牛肉背最长肌为原料，主要研究了牦牛组织蛋白酶L、B、H的提取分离过程及性质，牦牛肉成熟过程中组织蛋白酶L、B、H酶活力、肉品质及微观结构的变化，通过相关性分析阐明了组织蛋白酶L、B、H在成熟过程中的作用，为提高牦牛肉的品质、充分利用牦牛肉资源提供理论依据。
本研究包括四部分，具体内容和结果如下：
1.牦牛肉组织蛋白酶L、B及H的制备
以牦牛肉作为试验原料，依次采用缓冲液提取、盐析、透析、阴离子交换层析、分子筛层析等方法，最终纯化得到了三种分别具有Z-Phe-Arg-AMC、Z-Arg-Arg-AMC、L-Arg-AMC水解活性的组织蛋白酶，分别是组织蛋白酶L、B、H，收率分别为5.51、11.51、9.82%，纯化倍数分别为6.05、13.25、15.67fold，比活分别为1.15、0.53、0.47units/mg，总活分别为20.19、7.91、5.58U，分子量分别为36、35、27kDa。
2.牦牛肉组织蛋白酶L、B及H性质的研究
牦牛肉组织蛋白酶L、B、H的最适温度分别为50、45、50℃，最适pH分别为5.5、6.0、6.5。通过抑制剂、激活剂及金属离子对组织蛋白酶L、B、H酶活力的影响判断，三种蛋白酶都属于半胱氨酸类蛋白酶，而且活性中心都含有-SH。组织蛋白酶L、B、H的最适底物分别为Z-Phe-Arg-AMC、Z-Arg-Arg-AMC、L-Arg-AMC。组织蛋白酶L和B降解肌原纤维的最适pH为5.0~5.7，组织蛋白酶H降解肌原纤维的最适pH为4.0。
3.牦牛肉宰后成熟过程中组织蛋白酶L、B、H对肉品质变化的影响
牦牛肉成熟过程中组织蛋白酶L、B、H的酶活力随成熟时间的延长而增大，且组织蛋白酶L的活力高于组织蛋白酶B和H。成熟过程中，牦牛肉的pH值、糖原都发生了显著变化(P＜0.05)，而且牦牛肉的这些变化主要集中在宰后0~3d。成熟对牦牛肉的加压损失、蒸煮损失、剪切力、色度及质构都有影响，而对肉质最直接的影响是改善了其嫩度。通过相关性分析得出，成熟过程中组织蛋白酶L与pH呈显著负相关(P＜0.05)、组织蛋白酶B与pH呈极显著正相关(P＜0.01)，组织蛋白酶L、B、H与糖原均呈极显著负相关(P＜0.01)。
4.牦牛肉宰后成熟过程中组织蛋白酶L、B、H对微观结构变化的影响
牦牛肉宰后成熟过程中，MFI随成熟时间的延长而增大，在成熟前3d上升的速率和幅度较大；通过透射电镜观察发现，成熟8d牦牛肉肌原纤维结构完全被破坏，Z线断裂，M线完全消失；全肌肉蛋白电泳结果表明：成熟导致肌肉中的蛋白发生不同程度的降解，降解最明显的是肌钙蛋白；成熟可以使牦牛肉的肌纤维直径变小，从而提高了牦牛肉的嫩度。将MFI、肌纤维直径与组织蛋白酶L、B、H酶活力进行相关性分析，得出：组织蛋白酶L、B、H与MFI均呈极显著正相关(P＜0.01)，组织蛋白酶L、B、H与肌纤维直径均呈极显著负相关(P＜0.01)，肌纤维直径与MFI也呈极显著负相关(P＜0.01)。
综上所述，在宰后成熟过程中，牦牛肉的肉品质指标及微观结构均发生了变化，而这些变化导致的最直接的影响就是改善了牦牛肉的嫩度。根据组织蛋白酶L、B、H的活力、对肌原纤维的降解特性及其与肉品质、微观结构指标的相关性分析得出：组织蛋白酶H在成熟过程中几乎不起作用，组织蛋白酶L在成熟过程中的作用比B大。
It is well known that tenderness is one of the meat quality attributes consideredto be a determining factor in the acceptability and repurchasing intent of consumersand producers. Tenderness can be improved through postmortem storage atrefrigerated temperatures, a process called “conditioning”. The increase in tendernessduring postmortem ageing is due to the action of endogenous proteolytic enzymes.Three proteolytic systems have been implicated in the tenderization of meat:calcium-dependent proteases (calpains), the lysosomal proteases (cathepsins), andproteasomes. As yet, little information of cathepsins L, B and H was available in yakmeat. Accordingly, the research taking ten Datong yak Longissimus dorsi muscle(obtained randomly from Haiyan County, Qinghai province, China) as raw materials,the objectives of this work were to purify and characterize the endogenous cathepsinsL, B and H, observed the cathepsins L, B and H activity, meat quality andmicrostructure changes during postmortem ageing, through the correlation analysis toclarify the role of cathepsins L, B and H during postmortem ageing. This studyprovides further insights into the tenderization process of yak meat, which mayultimately be used for the advantageous manipulation of the process.
Cathepsins L, B and H from yak muscle were purified to6.05,13.25,15.67foldrespectively by buffer extract, ammonium sulfate fractionation and a series ofchromatographic separations. The specific activity of purified enzyme were1.15,0.53,0.47units/mg, yield coefficient were5.51,11.51,9.82%, total activity were20.19,7.91,5.58U and the molecular mass were36,35,27kDa as determined bySDS-PAGE.
The optimum temperature of cathepsins L, B, H were50℃,45℃and50℃respectively, optimum pH were5.5,6.0,6.5. Through observed the effect of inhibitors,activator and metal ions, suggested that the purified cathepsins belongs to cysteineproteinase family and the activity centers containing-SH. The optimum substrate ofcathepsins L, B and H were Z-Phe-Arg-AMC, Z-Arg-Arg-AMC, and L-Arg-AMCrespectively. The optimum pH of cathepsins L and B degradation myofibril were5.0~5.7, the optimum pH of cathepsin H was4.0.
Cathepsins L, B and H activities were at a minimum at the initial postmortemand increased significantly with time postmortem (P <0.05), and cathepsin L was detected in more important amounts than cathepsins B and H. pH, glycogen changedsignificantly (P <0.05), and these changes was mainly concentrated in the0~3dduring postmortem ageing. Ageing have impact on pressuring losses, cooking losses,shear force, color and texture parameters, and the most direct impact on meat is toimprove its tenderness. Correlation analysis showed, the cathepsins L and pH havesignificant correlation (P <0.05), cathepsin B and pH have highly significantcorrelation (P <0.01), and cathepsins L, B, H and glycogen was a significant negativecorrelation (P <0.01).
The MFI increased with time postmortem and relative to at-death values, anincrease (P <0.05) in MFI was detected during the first3d postmortem. Observed bytransmission electron microscope, we found that after8d ageing the muscle fibrilsstructure was completely disrupted, Z line rupture, M line disappear completely. Themuscle protein electrophoresis showed that ageing can lead to protein degradation ofthe different degree, the most obvious was Troponin. Ageing can make the yak musclefiber diameter decreases, thus improving the yak meat tenderness. Accordingcorrelation analysis with MFI, muscle fiber diameter and cathepsins L, B, H enzymeactivity, we concluded that cathepsins L, B, H and MFI have very significant positivecorrelated (P <0.01), cathepsins L, B, H and muscle fiber diameter have verynegatively significantly correlated (P <0.01), the muscle fiber diameter and MFIhave significant negative correlation (P <0.01).
In conclusion, meat quality and microstructure indexes also changed duringageing, these changes lead to the most direct impact was improved the tenderness ofyak meat. Accordance with the cathepsins L, B, H activity, myofibril degradationcharacteristic and the correlations between cathepsins L, B, H activity and meatquality, microstructure index, we found that cathepsin H had little effect on ageingprocess, the role of cathepsin L is greater than B.
本研究包括四部分，具体内容和结果如下：
1.牦牛肉组织蛋白酶L、B及H的制备
以牦牛肉作为试验原料，依次采用缓冲液提取、盐析、透析、阴离子交换层析、分子筛层析等方法，最终纯化得到了三种分别具有Z-Phe-Arg-AMC、Z-Arg-Arg-AMC、L-Arg-AMC水解活性的组织蛋白酶，分别是组织蛋白酶L、B、H，收率分别为5.51、11.51、9.82%，纯化倍数分别为6.05、13.25、15.67fold，比活分别为1.15、0.53、0.47units/mg，总活分别为20.19、7.91、5.58U，分子量分别为36、35、27kDa。
2.牦牛肉组织蛋白酶L、B及H性质的研究
牦牛肉组织蛋白酶L、B、H的最适温度分别为50、45、50℃，最适pH分别为5.5、6.0、6.5。通过抑制剂、激活剂及金属离子对组织蛋白酶L、B、H酶活力的影响判断，三种蛋白酶都属于半胱氨酸类蛋白酶，而且活性中心都含有-SH。组织蛋白酶L、B、H的最适底物分别为Z-Phe-Arg-AMC、Z-Arg-Arg-AMC、L-Arg-AMC。组织蛋白酶L和B降解肌原纤维的最适pH为5.0~5.7，组织蛋白酶H降解肌原纤维的最适pH为4.0。
3.牦牛肉宰后成熟过程中组织蛋白酶L、B、H对肉品质变化的影响
牦牛肉成熟过程中组织蛋白酶L、B、H的酶活力随成熟时间的延长而增大，且组织蛋白酶L的活力高于组织蛋白酶B和H。成熟过程中，牦牛肉的pH值、糖原都发生了显著变化(P＜0.05)，而且牦牛肉的这些变化主要集中在宰后0~3d。成熟对牦牛肉的加压损失、蒸煮损失、剪切力、色度及质构都有影响，而对肉质最直接的影响是改善了其嫩度。通过相关性分析得出，成熟过程中组织蛋白酶L与pH呈显著负相关(P＜0.05)、组织蛋白酶B与pH呈极显著正相关(P＜0.01)，组织蛋白酶L、B、H与糖原均呈极显著负相关(P＜0.01)。
4.牦牛肉宰后成熟过程中组织蛋白酶L、B、H对微观结构变化的影响
牦牛肉宰后成熟过程中，MFI随成熟时间的延长而增大，在成熟前3d上升的速率和幅度较大；通过透射电镜观察发现，成熟8d牦牛肉肌原纤维结构完全被破坏，Z线断裂，M线完全消失；全肌肉蛋白电泳结果表明：成熟导致肌肉中的蛋白发生不同程度的降解，降解最明显的是肌钙蛋白；成熟可以使牦牛肉的肌纤维直径变小，从而提高了牦牛肉的嫩度。将MFI、肌纤维直径与组织蛋白酶L、B、H酶活力进行相关性分析，得出：组织蛋白酶L、B、H与MFI均呈极显著正相关(P＜0.01)，组织蛋白酶L、B、H与肌纤维直径均呈极显著负相关(P＜0.01)，肌纤维直径与MFI也呈极显著负相关(P＜0.01)。
综上所述，在宰后成熟过程中，牦牛肉的肉品质指标及微观结构均发生了变化，而这些变化导致的最直接的影响就是改善了牦牛肉的嫩度。根据组织蛋白酶L、B、H的活力、对肌原纤维的降解特性及其与肉品质、微观结构指标的相关性分析得出：组织蛋白酶H在成熟过程中几乎不起作用，组织蛋白酶L在成熟过程中的作用比B大。
It is well known that tenderness is one of the meat quality attributes consideredto be a determining factor in the acceptability and repurchasing intent of consumersand producers. Tenderness can be improved through postmortem storage atrefrigerated temperatures, a process called “conditioning”. The increase in tendernessduring postmortem ageing is due to the action of endogenous proteolytic enzymes.Three proteolytic systems have been implicated in the tenderization of meat:calcium-dependent proteases (calpains), the lysosomal proteases (cathepsins), andproteasomes. As yet, little information of cathepsins L, B and H was available in yakmeat. Accordingly, the research taking ten Datong yak Longissimus dorsi muscle(obtained randomly from Haiyan County, Qinghai province, China) as raw materials,the objectives of this work were to purify and characterize the endogenous cathepsinsL, B and H, observed the cathepsins L, B and H activity, meat quality andmicrostructure changes during postmortem ageing, through the correlation analysis toclarify the role of cathepsins L, B and H during postmortem ageing. This studyprovides further insights into the tenderization process of yak meat, which mayultimately be used for the advantageous manipulation of the process.
Cathepsins L, B and H from yak muscle were purified to6.05,13.25,15.67foldrespectively by buffer extract, ammonium sulfate fractionation and a series ofchromatographic separations. The specific activity of purified enzyme were1.15,0.53,0.47units/mg, yield coefficient were5.51,11.51,9.82%, total activity were20.19,7.91,5.58U and the molecular mass were36,35,27kDa as determined bySDS-PAGE.
The optimum temperature of cathepsins L, B, H were50℃,45℃and50℃respectively, optimum pH were5.5,6.0,6.5. Through observed the effect of inhibitors,activator and metal ions, suggested that the purified cathepsins belongs to cysteineproteinase family and the activity centers containing-SH. The optimum substrate ofcathepsins L, B and H were Z-Phe-Arg-AMC, Z-Arg-Arg-AMC, and L-Arg-AMCrespectively. The optimum pH of cathepsins L and B degradation myofibril were5.0~5.7, the optimum pH of cathepsin H was4.0.
Cathepsins L, B and H activities were at a minimum at the initial postmortemand increased significantly with time postmortem (P <0.05), and cathepsin L was detected in more important amounts than cathepsins B and H. pH, glycogen changedsignificantly (P <0.05), and these changes was mainly concentrated in the0~3dduring postmortem ageing. Ageing have impact on pressuring losses, cooking losses,shear force, color and texture parameters, and the most direct impact on meat is toimprove its tenderness. Correlation analysis showed, the cathepsins L and pH havesignificant correlation (P <0.05), cathepsin B and pH have highly significantcorrelation (P <0.01), and cathepsins L, B, H and glycogen was a significant negativecorrelation (P <0.01).
The MFI increased with time postmortem and relative to at-death values, anincrease (P <0.05) in MFI was detected during the first3d postmortem. Observed bytransmission electron microscope, we found that after8d ageing the muscle fibrilsstructure was completely disrupted, Z line rupture, M line disappear completely. Themuscle protein electrophoresis showed that ageing can lead to protein degradation ofthe different degree, the most obvious was Troponin. Ageing can make the yak musclefiber diameter decreases, thus improving the yak meat tenderness. Accordingcorrelation analysis with MFI, muscle fiber diameter and cathepsins L, B, H enzymeactivity, we concluded that cathepsins L, B, H and MFI have very significant positivecorrelated (P <0.01), cathepsins L, B, H and muscle fiber diameter have verynegatively significantly correlated (P <0.01), the muscle fiber diameter and MFIhave significant negative correlation (P <0.01).
In conclusion, meat quality and microstructure indexes also changed duringageing, these changes lead to the most direct impact was improved the tenderness ofyak meat. Accordance with the cathepsins L, B, H activity, myofibril degradationcharacteristic and the correlations between cathepsins L, B, H activity and meatquality, microstructure index, we found that cathepsin H had little effect on ageingprocess, the role of cathepsin L is greater than B.
引文
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