巴西红耳龟红细胞血影蛋白质组学及对其低温缺氧抗性机理的研究
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摘要
龟类在动物界中属于脊索动物门、脊椎动物亚门、爬行纲、龟鳖亚纲、龟鳖目。龟类在进化史上属于活化石类的生物,以长寿而闻名,具有超强的耐低温和耐缺氧特性。耐低温耐缺氧机理研究具有非常重要的意义,例如在细胞冷冻过程中防止细胞被冰晶损伤,在器官移植手术中,离体器官长期保存而不会出现缺血/缺氧而造成的衰竭,保证再充血再充氧这样的再灌注过程中不被大量产生的自由基损伤,提高被远距离运送的重伤员的成活率等,甚至还有些仿生学科学家希望通过将人体通过类似的低温低氧“窖藏”的方式来延长生命。
龟类具有一些非常特殊的身体结构,其红细胞尤为特殊。龟类红细胞像大多数低等脊椎动物的红细胞一样为椭圆形,体积大且有核,不同之处在于成熟的龟类红细胞不含有线粒体。有的研究者指出龟类红细胞在进化史上与哺乳动物最接近,所以,可以把龟的红细胞定位于低等脊椎动物与高等脊椎动物红细胞之间。但是到目前为止,科学界对龟类红细胞的组成,尤其是对细胞保护,信号转导,物质运输有重要意义的细胞膜的组成知之甚少。如此特殊的红细胞结构及其与龟类超强的抗缺氧抗应激耐力的关系是本研究探讨的关键科学问题。本文以蛋白质组学及相关技术对巴西红耳龟血影的全蛋白质组学进行了研究,并发现了一些与巴西红耳龟低温缺氧耐受直接相关的蛋白,进而研究了其在巴西红耳龟低温缺氧过程中可能发挥的作用及作用机理。
首先,本文以巴西红耳龟(Red-eared turtle, Trachemys scripta)为研究材料,着眼于如何获得高纯度的巴西红耳龟红细胞血影,由于巴西红耳龟红细胞既有类似体细胞的复杂性,其细胞膜又具有哺乳动物红细胞的脆弱性,以至于分离体细胞的剧烈细胞破碎方法和分离哺乳动物红细胞的相对温和的低渗裂解法都不适用于分离龟类的红细胞。所以,纯化其红细胞血影是一项挑战。在本研究之前,未发现有详细的关于如何分离纯化巴西红耳龟红细胞血影的方法。本实验室在已有的研究报道的基础上通过反复摸索,最终总结出了一套有效的分离方法,通过富集红细胞,低渗裂解,具特殊针头注射器的吹打,低速匀浆以及差速离心的方法,我们最终获得了满足蛋白质组学鉴定要求的血影。并通过4种不同的样品前处理与不同的质谱鉴定方法相结合,介绍了一套有效的巴西红耳龟红细胞血影的蛋白质组学分析方法,共鉴定得到169个非冗余蛋白质并对其中有意义的蛋白质进行了深入的探讨,发现了一些与巴西红耳龟超强耐力相关的蛋白,如微管蛋白,热休克蛋白,以及定位在“红细胞膜上”的与产能直接相关的蛋白——F型ATP合酶亚单位。红细胞“非线粒体微生物样供能”为巴西红耳龟的抗逆生存机理研究打开了一个全新的视野。
其次,我们对巴西红耳龟进行了一周低温潜水处理,测定了一些与能量代谢相关的关键生化指标,发现了巴西红耳龟在短期低温潜水期间无氧糖酵解与糖异生同时进行的能量代谢方式。对短期低温潜水期间巴西红耳龟体内肝糖原和肌糖原储备没有被消耗而血糖升高的现象作出了独特的阐释。
最后,我们结合短期低温潜水期间能量代谢的特点以及前人的研究基础,进行了血液体外ATP剥夺实验,证实了巴西红耳龟红细胞具有向细胞外产生ATP的特性。可能的产能机理是红细胞利用细胞内比细胞外高出约5倍的氢离子推动细胞膜上的ATP合酶合成ATP。并在此基础上提出了巴西红耳龟短期低温潜水期间的能量代谢模型假说。
总之,本研究从巴西红耳龟血影的分离纯化到血影蛋白的高效酶切,再到质谱鉴定,生物信息学分析,系统地提出了一套针对巴西红耳龟甚至是有核红细胞血影的组学分析方法。同时,对巴西红耳龟短期低温缺氧能量代谢机理进行了探讨,发现巴西红耳龟红细胞除了携带氧的作用外,还可能具有供能作用。本课题的研究发现为今后更加深入地研究巴西红耳龟以至整个龟类的能量代谢机制和低温缺氧应对机制提供了全新的有重要进化生物学价值和临床医学意义的实验数据和理论假说。
Turtles in the animal kingdom belong to the chordates, vertebrate subphylum, reptilia, turtle subclass, chelonia. Turtles are living fossil creatures during the evolutionary history. Turtles are famous as longevity and its super-tolerance to low-temperature and hypoxia are impressing. Scientists have fascinated at the mechanism of low-temperature and hypoxia tolerance for many years. The research on low-temperature/hypoxia tolerance mechanism has very important significance. Such as prevent the cells be damaged by ice crystal during freezing, In vitro, organ transplants fields, preserve isolated organ long-term without failure, which caused by ischemia/hypoxia, avoid the free-radical damage during the process of re-oxygenation or ischemia and reperfusion, improve the survival rate of injured personnel who have to be transported over a long distance to medical care, etc. And even some scientists hope to prolong lifespan similar to low-temperature/hypoxia cellaring.
The turtles have some very specific body composition, such as their erythrocytes. Turtle erythrocyte has the same oval-shaped, huge, and nuclear as the other red blood cell (RBC) of low vertebrates, however, mature trutle RBCs does not contain mitochondria, which is a common existence in most of the low vertebrate RBCs. Some researchers consider that the turtle erythrocyte is closest with mammals RBC during evolutionary history. Erythrocyte of turtle between the RBCs of low vertebrates and higher vertebrates. However, the researchers still know very little about the components of turtle erythrocyte. Especially about cell membrane which plays an important role in cell protection, signal transduction, material transport, etc. in cell. The special structure of turtle erythrocyte may have direct correlation with the super endurance. Red-eared turtles were used to analyze the erythrocyte ghost proteomics in this study. We hope to partially explain the mechanism of low-temperature and hypoxia tolerance of turtle.
First, we focused on how to obtain high-purity red-eared turtle erythrocyte ghost (TEG). As nucleated cells, turtle erythrocytes have a complexity of somatic cell; on the other hand, as RBCs, its plasma membrane has the fragility as mature human RBCs. Purification method of somatic cell and mature mammalian erythrocyte ghosts is unsuitable for the TEG. Therefore, how to purify TEG is a huge challenge. Before this study, we did not find any detailed information on how to separate and purify TEG. Based on existing knowledge and combined with our repeated experiments, eventually we summarized an effective separation method for the preparation of TEG. Namely, after hypotonic lysis with specific buffer, forcing through a syringe with specially shaped needle, low speed homogenizes and differential centrifugation, highly purified TEG fractions were separated effectively, which is a key to successful TEG proteomics. The TEG proteins were digested and dissociated in4different methods combined with different mass spectrum analysis; a total of169TEG proteins was identified. After further bioinformatic analyzing of these identified proteins, tubulins, heat shock proteins were identified in the TEG, which provided important insights into the low-temperature and hypoxia tolerance of Trachemys scripta. We also identified membrane located F-type ATP synthase subunits, which closely related with the production of ATP. The proteomics analysis of TEG opened a new direction of super-tolerance research of red-eared turtle.
Secondly, we studied the important biochemical indicators energy metabolism of red-eared turtle by short-term low-temperature and diving exposure. We found anaerobic glycolysis and gluconeogenesis simultaneously during this short term low-temperature/hypoxic exposure. This finding gave a good explanation for why liver glycogen and muscle glycogen reserved but blood glucose elevated during a shot-term low-temperature and diving exposure.
Finally, we combined the characteristics of the energy metabolism of short-term low temperature and diving exposure, blood ATP deprivation experiment in vitro and previous studies; we found that red-eared turtle red blood cells can generate ATP to the extracellular. The possible mechanism was high concentration of H+promoting the RBC plasma membrane located ATP synthase generated ATP. Meanwhile, we propose a model of RBCs energy supply for short-term low-temperature/hypoxia exposure.
Therefore, in this study, an effective isolation method to prepare TEG was developed in combination with predissociation, enzymolysis and CapLC-MS/MS identification. A data set of TEG proteins were provided, which allows for more comprehensive characterization of TEG. Furthermore, we studied the energy metabolism mechanism of low-temperature/hypoxia red-eared turtle. We opened a new research field for understanding the super-tolerance of turtles when they face severe environment.
龟类具有一些非常特殊的身体结构,其红细胞尤为特殊。龟类红细胞像大多数低等脊椎动物的红细胞一样为椭圆形,体积大且有核,不同之处在于成熟的龟类红细胞不含有线粒体。有的研究者指出龟类红细胞在进化史上与哺乳动物最接近,所以,可以把龟的红细胞定位于低等脊椎动物与高等脊椎动物红细胞之间。但是到目前为止,科学界对龟类红细胞的组成,尤其是对细胞保护,信号转导,物质运输有重要意义的细胞膜的组成知之甚少。如此特殊的红细胞结构及其与龟类超强的抗缺氧抗应激耐力的关系是本研究探讨的关键科学问题。本文以蛋白质组学及相关技术对巴西红耳龟血影的全蛋白质组学进行了研究,并发现了一些与巴西红耳龟低温缺氧耐受直接相关的蛋白,进而研究了其在巴西红耳龟低温缺氧过程中可能发挥的作用及作用机理。
首先,本文以巴西红耳龟(Red-eared turtle, Trachemys scripta)为研究材料,着眼于如何获得高纯度的巴西红耳龟红细胞血影,由于巴西红耳龟红细胞既有类似体细胞的复杂性,其细胞膜又具有哺乳动物红细胞的脆弱性,以至于分离体细胞的剧烈细胞破碎方法和分离哺乳动物红细胞的相对温和的低渗裂解法都不适用于分离龟类的红细胞。所以,纯化其红细胞血影是一项挑战。在本研究之前,未发现有详细的关于如何分离纯化巴西红耳龟红细胞血影的方法。本实验室在已有的研究报道的基础上通过反复摸索,最终总结出了一套有效的分离方法,通过富集红细胞,低渗裂解,具特殊针头注射器的吹打,低速匀浆以及差速离心的方法,我们最终获得了满足蛋白质组学鉴定要求的血影。并通过4种不同的样品前处理与不同的质谱鉴定方法相结合,介绍了一套有效的巴西红耳龟红细胞血影的蛋白质组学分析方法,共鉴定得到169个非冗余蛋白质并对其中有意义的蛋白质进行了深入的探讨,发现了一些与巴西红耳龟超强耐力相关的蛋白,如微管蛋白,热休克蛋白,以及定位在“红细胞膜上”的与产能直接相关的蛋白——F型ATP合酶亚单位。红细胞“非线粒体微生物样供能”为巴西红耳龟的抗逆生存机理研究打开了一个全新的视野。
其次,我们对巴西红耳龟进行了一周低温潜水处理,测定了一些与能量代谢相关的关键生化指标,发现了巴西红耳龟在短期低温潜水期间无氧糖酵解与糖异生同时进行的能量代谢方式。对短期低温潜水期间巴西红耳龟体内肝糖原和肌糖原储备没有被消耗而血糖升高的现象作出了独特的阐释。
最后,我们结合短期低温潜水期间能量代谢的特点以及前人的研究基础,进行了血液体外ATP剥夺实验,证实了巴西红耳龟红细胞具有向细胞外产生ATP的特性。可能的产能机理是红细胞利用细胞内比细胞外高出约5倍的氢离子推动细胞膜上的ATP合酶合成ATP。并在此基础上提出了巴西红耳龟短期低温潜水期间的能量代谢模型假说。
总之,本研究从巴西红耳龟血影的分离纯化到血影蛋白的高效酶切,再到质谱鉴定,生物信息学分析,系统地提出了一套针对巴西红耳龟甚至是有核红细胞血影的组学分析方法。同时,对巴西红耳龟短期低温缺氧能量代谢机理进行了探讨,发现巴西红耳龟红细胞除了携带氧的作用外,还可能具有供能作用。本课题的研究发现为今后更加深入地研究巴西红耳龟以至整个龟类的能量代谢机制和低温缺氧应对机制提供了全新的有重要进化生物学价值和临床医学意义的实验数据和理论假说。
Turtles in the animal kingdom belong to the chordates, vertebrate subphylum, reptilia, turtle subclass, chelonia. Turtles are living fossil creatures during the evolutionary history. Turtles are famous as longevity and its super-tolerance to low-temperature and hypoxia are impressing. Scientists have fascinated at the mechanism of low-temperature and hypoxia tolerance for many years. The research on low-temperature/hypoxia tolerance mechanism has very important significance. Such as prevent the cells be damaged by ice crystal during freezing, In vitro, organ transplants fields, preserve isolated organ long-term without failure, which caused by ischemia/hypoxia, avoid the free-radical damage during the process of re-oxygenation or ischemia and reperfusion, improve the survival rate of injured personnel who have to be transported over a long distance to medical care, etc. And even some scientists hope to prolong lifespan similar to low-temperature/hypoxia cellaring.
The turtles have some very specific body composition, such as their erythrocytes. Turtle erythrocyte has the same oval-shaped, huge, and nuclear as the other red blood cell (RBC) of low vertebrates, however, mature trutle RBCs does not contain mitochondria, which is a common existence in most of the low vertebrate RBCs. Some researchers consider that the turtle erythrocyte is closest with mammals RBC during evolutionary history. Erythrocyte of turtle between the RBCs of low vertebrates and higher vertebrates. However, the researchers still know very little about the components of turtle erythrocyte. Especially about cell membrane which plays an important role in cell protection, signal transduction, material transport, etc. in cell. The special structure of turtle erythrocyte may have direct correlation with the super endurance. Red-eared turtles were used to analyze the erythrocyte ghost proteomics in this study. We hope to partially explain the mechanism of low-temperature and hypoxia tolerance of turtle.
First, we focused on how to obtain high-purity red-eared turtle erythrocyte ghost (TEG). As nucleated cells, turtle erythrocytes have a complexity of somatic cell; on the other hand, as RBCs, its plasma membrane has the fragility as mature human RBCs. Purification method of somatic cell and mature mammalian erythrocyte ghosts is unsuitable for the TEG. Therefore, how to purify TEG is a huge challenge. Before this study, we did not find any detailed information on how to separate and purify TEG. Based on existing knowledge and combined with our repeated experiments, eventually we summarized an effective separation method for the preparation of TEG. Namely, after hypotonic lysis with specific buffer, forcing through a syringe with specially shaped needle, low speed homogenizes and differential centrifugation, highly purified TEG fractions were separated effectively, which is a key to successful TEG proteomics. The TEG proteins were digested and dissociated in4different methods combined with different mass spectrum analysis; a total of169TEG proteins was identified. After further bioinformatic analyzing of these identified proteins, tubulins, heat shock proteins were identified in the TEG, which provided important insights into the low-temperature and hypoxia tolerance of Trachemys scripta. We also identified membrane located F-type ATP synthase subunits, which closely related with the production of ATP. The proteomics analysis of TEG opened a new direction of super-tolerance research of red-eared turtle.
Secondly, we studied the important biochemical indicators energy metabolism of red-eared turtle by short-term low-temperature and diving exposure. We found anaerobic glycolysis and gluconeogenesis simultaneously during this short term low-temperature/hypoxic exposure. This finding gave a good explanation for why liver glycogen and muscle glycogen reserved but blood glucose elevated during a shot-term low-temperature and diving exposure.
Finally, we combined the characteristics of the energy metabolism of short-term low temperature and diving exposure, blood ATP deprivation experiment in vitro and previous studies; we found that red-eared turtle red blood cells can generate ATP to the extracellular. The possible mechanism was high concentration of H+promoting the RBC plasma membrane located ATP synthase generated ATP. Meanwhile, we propose a model of RBCs energy supply for short-term low-temperature/hypoxia exposure.
Therefore, in this study, an effective isolation method to prepare TEG was developed in combination with predissociation, enzymolysis and CapLC-MS/MS identification. A data set of TEG proteins were provided, which allows for more comprehensive characterization of TEG. Furthermore, we studied the energy metabolism mechanism of low-temperature/hypoxia red-eared turtle. We opened a new research field for understanding the super-tolerance of turtles when they face severe environment.
引文
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