罗非鱼PPARγ在脂代谢中的调节作用及其转录活性
发布时间：2018-08-09 12:54
【摘要】：尼罗罗非鱼(Oreochromis niloticus)生长性能好且抗逆性强,已成为世界性养殖品种。作为一种重要的模式物种,其基因组测序已经完成,有利于以其为模型开展鱼类营养生理研究。过氧化物体增殖物激活受体γ(Peroxisome proliferator activated receptor gamma, PPARy)是动物脂类代谢关键调节器,它广泛存在于脊椎动物中。然而,目前尚缺乏有关鱼类PPARy生理功能、分子结构及转录活性等方面的研究。本研究以尼罗罗非鱼为模型,采用分子克隆、细胞培养、荧光定量PCR、组织学分析、蛋白印迹、转录组学、双荧光素报道系统、启动子捕获和代谢物含量测定等手段针对PPARy进行以下四方面的研究：1)罗非鱼PPARy在调节脂质内稳态中的作用；2)罗非鱼PPARy结构分析及转录活性研究；3)哺乳动物PPARy激活剂Rosiglitazone和抑制剂GW9662对罗非鱼脂代谢的影响,和4)PPARy靶基因ACOX1的克隆及其营养调控。论文的主要结果与结论如下1.尼罗罗非鱼PPARy在脂质内稳态中的调节作用自然选择不仅赋予了动物在脂类物质丰富时贮存脂类的能力,同时也赋予了其在外界脂类摄入不足时进行体内脂类合成的能力。然而鱼类是如何利用自身的脂代谢系统来应对低脂食物和高脂食物的不同营养条件仍未得到很好的阐明。本研究通过高脂饲料和低脂饲料投喂法来探究尼罗罗非鱼在脂类摄入过量和不足这两种情况下是如何维持脂类内稳态的,以及PPARy在这一过程中扮演了何种角色。实验结果显示,持续10周投喂脂肪含量分别为1%,7%和13%三种饲料后,三组之间的生长率(Growth rate),肝体比(Hepatic somatic index),血清、肝脏、肌肉和脂肪组织的甘油三酯含量均无显著差异,但是体脂肪含量和脂体比随着饲料中脂肪水平的增加而增加。荧光定量PCR、转录组学、蛋白免疫印迹等实验结果表明,肝脏是应对低脂投喂的主要器官,主要表现为糖酵解速率增加和脂肪酸从头合成能力增强；在高脂条件下,脂肪组织通过吸收更多游离脂肪酸并增加脂肪水解,进而激活PPARγ,被激活的PPARγ进一步促进脂肪细胞增殖,从而使脂肪细胞能在不变大的情况下贮存更多甘油三酯,此研究结果表明罗非鱼PPARy有着与哺乳动物相似的功能。此研究首次较为系统的阐明了鱼类应对脂肪摄入不足和过量时的生理调控机制,丰富了我们对鱼类生理学的认识。2.尼罗罗非鱼PPARy结构分析及转录活性研究过氧化物酶体增殖物激活受体y (Peroxisome proliferator activated receptor gamma, PPARy)是动物脂类代谢的关键调节器,前节表明尼罗罗非鱼PPARy参与调节罗非鱼脂肪细胞增殖,分子结构决定了其生理功能,因此在此基础上,我们进一步对PPARy的分子结构和转录活性进行了分析。本研究首先获得了尼罗罗非鱼PPARy全长(NtPPARy),并通过构建NtPPARy和NtRXRa表达载体,及FABP4启动子驱动的报道载体在HEK-293细胞系中开展了转录活性研究。结果发现：NtPPARy存在两种转录本,二者在5’-非编码区存在差异,与哺乳动物比较后发现,NtPPARy的LBD (Ligand binding domain)多了39个氨基酸残基,使得罗非鱼LBD区比哺乳动物多3个alpha螺旋结构。组织表达模式结果发现两种转录本在11种组织中呈现出不同的分布模式,但两者均在肝脏、肠和肾脏中高表达。转录活性研究结果显示NtPPARy与NtRXRa一起参与尼罗罗非鱼FABP4基因的转录调节,与人类中的转录调节方式相似。总之,PPARy的DBD (DNA binding domain)高度保守,但LBD保守性相对较弱,而且罗非鱼PPARy与RXRa一起参与诸如FABP4等靶基因的转录调节。在获得尼罗罗非鱼细胞系的基础上,本研究所构建的质粒系统将会是未来研究鱼类PPARy功能的有效工具。3.哺乳动物PPARy激活剂Rosiglitazone和抑制剂GW9662对罗非鱼脂代谢的影响功能获得(Gain of function)和功能缺失(Loss of function)是研究基因功能的有效策略,利用激活剂和抑制剂可以快速实现功能获得和功能缺失,因此我们尝试通过哺乳动物PPARy激活剂Rosiglitazone (Rosi)和抑制剂GW9662(GW9)来激活和抑制罗非鱼PPARy,并在此基础上探讨罗非鱼PPARy的生理功能。本研究首先设计二种脂肪水平的饲料,即4%(Standard diet, SD)和15%(High fat diet,HFD),每种水平下包括对照组,Rosi (15mg/kg)组,和GW9 (10mg/kg)组,共6组饲料,养殖6周后采样分析。结果发现HFD组的脂体比显著高于SD组,肝体比和肝脏中TG含量也高于SD组,表明高脂导致肝脏脂肪沉积过量；HFD组血清TG含量也显著高于SD组,表明脂质内稳态失衡,成功获得我们所需的脂肪肝模型。但无论是在低脂还是高脂背景下,Rosi和GW9的添加对罗非鱼生长和代谢指标均无显著影响。因此我们采用腹腔注射这一更为直接的给药方式进行二次实验,Rosiglitazone注射剂量为：0(DMSO)、0.6mg/kg、18mg/kg和54 mg/kg;GW9662注射剂量为：0(DMSO)、0.4mg/kg、12mg/kg和36 mg/kg,连续注射三天后采样分析。腹腔注射GW9662对血清中甘油三酯(Triglyceride, TG)、游离脂肪酸(Free fatty acid, FFA)和葡萄糖(Glucose)均无显著影响。注射Rosiglitazone对血糖无影响,但是显著下调了TG含量,上调了FFA含量。荧光定量PCR分析发现肝脏中脂肪酸合成,TG合成和VLDL关键蛋白基因表达量均下降,表明TG含量下降主要归功于肝脏脂质合成和分泌减少,FFA的升高则归功于肌肉中脂肪水解增强。但是Rosi没有影响PPARγ在脂肪组织、肝脏和肌肉中的表达,是否调节了PPARγ的转录后修饰,并影响了其蛋白活性从而引起相应的生理效应则需进一步研究。以上结果表明,短期Rosi注射能够促进罗非鱼肌肉脂肪水解。4. PPARγ靶基因ACOX1的克隆及其营养调控过氧化物酶体是合成多种脂类信号分子的场所。过氧化物酶体中的代谢活动受到PPARγ的调节,酯酰辅酶A氧化酶1(Acayl-CoA Oxidase 1, ACOX1)是过氧化物酶体β-氧化的第一限速酶,它参与酯酰辅酶A至2-顺式烯酰辅酶A的催化过程,而过氧化物酶体β-氧化过程为信号分子的合成提供底物。ACOX1是PPARγ的靶基因之一,因此阐明ACOX1的结构和功能对研究PPARγ如何调控过氧化物体信号分子的合成具有重要作用。本研究首次在尼罗罗非鱼中克隆获得了ACOX1基因,发现其具有两种转录本,分别命名为ACOXli1和ACOXli2,它们编码的蛋白均由661个氨基酸残基组成。两亚型的编码区由14个外显子组成。ACOXli1的89-193号位氨基酸由外显子E3b编码,而ACOXli2,的贝由E3a编码,同源比对分析发现ACOXli1的保守性明显高于ACOXli2,提示ACOXli1的功能可能比ACOXli2的功能更为重要。二者的mRNA组织分布模式表明：ACOXi1在肝中的表达量最高,其次为肾脏和脑；而ACOXli2在肾脏中表达量最高,其次为肝脏,两种亚型在白肌、鳃、脑和肾脏中的表达量呈显著差异,在白肌和肾脏中ACOXli2显著高于ACOXlil,而在脑和鳃中则相反；在饥饿36小时、饱食后1、3、8、24小时的营养状态研究中发现：两种亚型在肝中对营养状态做出了相同的反应模式,肾脏ACOXlil与肝中的反应模式趋同,而ACOXli2不受营养状态的影响。以上实验结果表明ACOXl基因两种亚型在脊椎动物中可能行使不同的功能。
[Abstract]:Oreochromis niloticus (Nile Tilapia) has good growth performance and strong resistance, and has become a world breed. As an important model species, its genome sequencing has been completed, and it is beneficial to carry out the study of fish nutrition physiology with its model. Peroxisome proliferator activated recept (Peroxisome proliferator activated recept) Or gamma, PPARy, a key regulator of animal lipid metabolism, is widely found in vertebrates. However, there is still a lack of research on the physiological function, molecular structure and transcriptional activity of fish PPARy. This study uses Nile tilapia as a model, using molecular cloning, cell culture, fluorescence quantitative PCR, histological analysis, and Western blot. Transcriptional studies, double fluorescein reporting system, promoter capture and metabolite content determination are used to study the following four aspects of PPARy: 1) the role of PPARy in regulating homeostasis of lipid; 2) PPARy structure analysis and transcriptional activity study of tilapia; 3) mammalian activator Rosiglitazone and inhibitor GW9662 The effect of lipid metabolism of tilapia, and 4) the cloning and nutrition regulation of the PPARy target gene ACOX1. The main results and conclusions of this paper are as follows: 1. the natural selection of PPARy in the homeostasis of lipid in Nile tilapia not only endows the energy of the animals when they are rich in lipid, but also endows them with the intake of lipids in the outside world. The ability to synthesize lipids in the body is carried out in the body. However, fish is not well elucidated on how to use its own lipid metabolism system to cope with the different nutritional conditions of low fat and high fat foods. In this study, the two cases of Nile tilapia were investigated by high fat feed and low fat feed. How to maintain the homeostasis in the lipid and what role PPARy plays in this process. The results showed that the growth rate (Growth rate) between the three groups (Hepatic somatic index), the liver body ratio (Hepatic somatic index), the serum, the liver, the muscle and the fat tissues were all content after 10 weeks of feeding of fat and three kinds of feed, respectively. There was no significant difference, but the body fat content and fat body ratio increased with the increase of fat level in the feed. The results of fluorescence quantitative PCR, transcriptional omics, and protein immunoblotting showed that the liver was the main organ to respond to low fat feeding, mainly the increase of glycolysis rate and the enhancement of adipose acid ab initio synthesis ability; in high fat condition. At the same time, the adipose tissue activates PPAR gamma by absorbing more free fatty acids and increasing the hydrolysis of fat, and then activates the activated PPAR gamma to further promote the proliferation of adipocytes, so that the fat cells can store more triglycerides in the same condition. The results show that the PPARy has a similar function to the mammal. The physiological regulation mechanism of fish in response to insufficient and excessive fat intake was systematically clarified, which enriched our understanding of fish physiology,.2. Nile tilapia PPARy structure analysis and transcriptional activity study, the peroxisome proliferator activation receptor y (Peroxisome proliferator activated receptor gamma, PPARy) is animal fat The key regulator of class metabolism, the previous section shows that Nile tilapia PPARy is involved in regulating the proliferation of tilapia adipocyte, and the molecular structure determines its physiological function. On this basis, we further analyzed the molecular structure and transcriptional activity of PPARy. First, the total length of PPARy (NtPPARy) of Nile tilapia was obtained and passed. The transcriptional activity of the NtPPARy and NtRXRa expression vectors and the FABP4 promoter driven report carrier in the HEK-293 cell line was studied. The results showed that there were two transcripts in NtPPARy, the two in the 5 '- non coding region, and the NtPPARy LBD (Ligand binding domain) more than 39 amino acid residues after comparison with the mammalian. The tissue expression pattern showed that the two transcripts showed different distribution patterns in the 11 tissues, but both were highly expressed in the liver, the intestines and the kidneys. The transcriptional activity study showed that NtPPARy and NtRXRa were involved in the transcription of the FABP4 gene of Nile tilapia (Nile Tilapia). The results showed that NtPPARy and NtRXRa were involved in the transcription of the FABP4 gene. In conclusion, the DBD (DNA binding domain) of PPARy is highly conservative, but the LBD conservatism is relatively weak, and the tilapia PPARy and RXRa participate in the transcriptional regulation of the target genes such as FABP4. On the basis of obtaining the Nile tilapia cell line, the plasmid system of this study will be the future. An effective tool for studying fish PPARy function.3. mammalian PPARy activator Rosiglitazone and inhibitor GW9662 on lipid metabolism of tilapia (Gain of function) and functional deletion (Loss of function) are effective strategies for the study of gene function. Functional acquisition and functional deficiency can be quickly realized by using activators and inhibitors. Therefore, we tried to activate and inhibit the PPARy of tilapia by mammalian PPARy activator Rosiglitazone (Rosi) and inhibitor GW9662 (GW9). On this basis, we explored the physiological functions of the tilapia PPARy. This study first designed two kinds of fat level feedstuff, namely, 4% (Standard diet, SD) and 15% (High fat), at each level. Including the control group, the Rosi (15mg/kg) group, and the GW9 (10mg/kg) group, a total of 6 groups of feed were collected and analyzed after 6 weeks of culture. The results showed that the lipid body ratio of the HFD group was significantly higher than that in the SD group. The liver body ratio and the TG content in the liver were also higher than those in the SD group, indicating that the high fat led to the excessive fat deposition in the liver, and the serum TG content of the HFD group was significantly higher than that in the SD group, indicating the homeostasis imbalance in the lipid. We successfully obtained the fatty liver model we needed. However, the addition of Rosi and GW9 had no significant effect on the growth and metabolism of tilapia in low fat or high lipid background. Therefore, two experiments were carried out by intraperitoneal injection of this more direct method. The amount of Rosiglitazone injection was 0 (DMSO), 0.6mg/kg, 18mg/kg and 5. 4 mg/kg; the amount of GW9662 injection was 0 (DMSO), 0.4mg/kg, 12mg/kg, and 36 mg/kg, and the injection of GW9662 on serum triglyceride (Triglyceride, TG), free fatty acids (Free fatty acid), and glucose had no significant effect on blood glucose. TG content increased the content of FFA. Fluorescence quantitative PCR analysis found that fatty acid synthesis in the liver, TG synthesis and VLDL key protein gene expression decreased, indicating that the decrease of TG content was mainly attributed to the decrease of liver lipid synthesis and secretion, and the increase of FFA was attributed to the enhancement of lipid hydrolysis in the muscles. But Rosi did not affect the PPAR gamma in adipose tissue. The expression in the liver and muscle, which regulates the post transcriptional modification of PPAR gamma, affects its protein activity and causes the corresponding physiological effects. The results suggest that short term Rosi injection can promote the cloning of the lipid hydrolysis of the.4. PPAR gamma target gene ACOX1 of the tilapia muscle and its nutritional regulation of peroxisomes. The metabolic activity in peroxisomes is regulated by PPAR gamma, and the ester acyl coenzyme A oxidase 1 (Acayl-CoA Oxidase 1, ACOX1) is the first rate limiting enzyme of peroxisome beta oxidation, and it participates in the catalytic process of ester acyl coenzyme A to 2- CIS enyl coenzyme A, and peroxisome beta oxidation process as a letter. The synthesis of molecule.ACOX1 is one of the target genes of PPAR gamma. Therefore, it is important to elucidate the structure and function of ACOX1 for the study of how PPAR gamma regulates the synthesis of signal molecules of peroxidation objects. In this study, the ACOX1 gene was cloned in Nile tilapia for the first time. It was found to have two transcripts, named ACOXli1, respectively. And ACOXli2, their encoded proteins are composed of 661 amino acid residues. The two subtype of the encoding region is encoded by the exon E3b, which consists of 14 exons, and the ACOXli2, the ACOXli2, is encoded by E3a. The homology analysis shows that the conservatism of ACOXli1 is significantly higher than that of ACOXli2, suggesting that ACOXli1's function may be more than ACOXli2. The mRNA distribution pattern of the two groups showed that the expression of ACOXi1 in the liver was the highest, followed by the kidney and the brain, and the expression of ACOXli2 in the kidney was the highest, followed by the liver. The expression of the two subtypes in the white muscles, gills, brain and kidneys was significantly different, and the ACOXli2 was significantly higher in the white muscles and kidneys than in the ACOXlil, while in the brain, the brain was significantly higher than that in the brain. In contrast to the gills, in the 36 hours of hunger, the nutritional status of 1,3,8,24 hours after full food was found: the two subtypes made the same response pattern in the liver, the kidney ACOXlil and the liver response patterns converged, and the ACOXli2 was not affected by the nutritional status. The experimental results showed that the two subtypes of the ACOXl gene were in the ridge. Different functions may be exercised in vertebroid animals.
【学位授予单位】：华东师范大学
【学位级别】：博士
【学位授予年份】：2016
【分类号】：Q953
，
本文编号：2174136
[Abstract]:Oreochromis niloticus (Nile Tilapia) has good growth performance and strong resistance, and has become a world breed. As an important model species, its genome sequencing has been completed, and it is beneficial to carry out the study of fish nutrition physiology with its model. Peroxisome proliferator activated recept (Peroxisome proliferator activated recept) Or gamma, PPARy, a key regulator of animal lipid metabolism, is widely found in vertebrates. However, there is still a lack of research on the physiological function, molecular structure and transcriptional activity of fish PPARy. This study uses Nile tilapia as a model, using molecular cloning, cell culture, fluorescence quantitative PCR, histological analysis, and Western blot. Transcriptional studies, double fluorescein reporting system, promoter capture and metabolite content determination are used to study the following four aspects of PPARy: 1) the role of PPARy in regulating homeostasis of lipid; 2) PPARy structure analysis and transcriptional activity study of tilapia; 3) mammalian activator Rosiglitazone and inhibitor GW9662 The effect of lipid metabolism of tilapia, and 4) the cloning and nutrition regulation of the PPARy target gene ACOX1. The main results and conclusions of this paper are as follows: 1. the natural selection of PPARy in the homeostasis of lipid in Nile tilapia not only endows the energy of the animals when they are rich in lipid, but also endows them with the intake of lipids in the outside world. The ability to synthesize lipids in the body is carried out in the body. However, fish is not well elucidated on how to use its own lipid metabolism system to cope with the different nutritional conditions of low fat and high fat foods. In this study, the two cases of Nile tilapia were investigated by high fat feed and low fat feed. How to maintain the homeostasis in the lipid and what role PPARy plays in this process. The results showed that the growth rate (Growth rate) between the three groups (Hepatic somatic index), the liver body ratio (Hepatic somatic index), the serum, the liver, the muscle and the fat tissues were all content after 10 weeks of feeding of fat and three kinds of feed, respectively. There was no significant difference, but the body fat content and fat body ratio increased with the increase of fat level in the feed. The results of fluorescence quantitative PCR, transcriptional omics, and protein immunoblotting showed that the liver was the main organ to respond to low fat feeding, mainly the increase of glycolysis rate and the enhancement of adipose acid ab initio synthesis ability; in high fat condition. At the same time, the adipose tissue activates PPAR gamma by absorbing more free fatty acids and increasing the hydrolysis of fat, and then activates the activated PPAR gamma to further promote the proliferation of adipocytes, so that the fat cells can store more triglycerides in the same condition. The results show that the PPARy has a similar function to the mammal. The physiological regulation mechanism of fish in response to insufficient and excessive fat intake was systematically clarified, which enriched our understanding of fish physiology,.2. Nile tilapia PPARy structure analysis and transcriptional activity study, the peroxisome proliferator activation receptor y (Peroxisome proliferator activated receptor gamma, PPARy) is animal fat The key regulator of class metabolism, the previous section shows that Nile tilapia PPARy is involved in regulating the proliferation of tilapia adipocyte, and the molecular structure determines its physiological function. On this basis, we further analyzed the molecular structure and transcriptional activity of PPARy. First, the total length of PPARy (NtPPARy) of Nile tilapia was obtained and passed. The transcriptional activity of the NtPPARy and NtRXRa expression vectors and the FABP4 promoter driven report carrier in the HEK-293 cell line was studied. The results showed that there were two transcripts in NtPPARy, the two in the 5 '- non coding region, and the NtPPARy LBD (Ligand binding domain) more than 39 amino acid residues after comparison with the mammalian. The tissue expression pattern showed that the two transcripts showed different distribution patterns in the 11 tissues, but both were highly expressed in the liver, the intestines and the kidneys. The transcriptional activity study showed that NtPPARy and NtRXRa were involved in the transcription of the FABP4 gene of Nile tilapia (Nile Tilapia). The results showed that NtPPARy and NtRXRa were involved in the transcription of the FABP4 gene. In conclusion, the DBD (DNA binding domain) of PPARy is highly conservative, but the LBD conservatism is relatively weak, and the tilapia PPARy and RXRa participate in the transcriptional regulation of the target genes such as FABP4. On the basis of obtaining the Nile tilapia cell line, the plasmid system of this study will be the future. An effective tool for studying fish PPARy function.3. mammalian PPARy activator Rosiglitazone and inhibitor GW9662 on lipid metabolism of tilapia (Gain of function) and functional deletion (Loss of function) are effective strategies for the study of gene function. Functional acquisition and functional deficiency can be quickly realized by using activators and inhibitors. Therefore, we tried to activate and inhibit the PPARy of tilapia by mammalian PPARy activator Rosiglitazone (Rosi) and inhibitor GW9662 (GW9). On this basis, we explored the physiological functions of the tilapia PPARy. This study first designed two kinds of fat level feedstuff, namely, 4% (Standard diet, SD) and 15% (High fat), at each level. Including the control group, the Rosi (15mg/kg) group, and the GW9 (10mg/kg) group, a total of 6 groups of feed were collected and analyzed after 6 weeks of culture. The results showed that the lipid body ratio of the HFD group was significantly higher than that in the SD group. The liver body ratio and the TG content in the liver were also higher than those in the SD group, indicating that the high fat led to the excessive fat deposition in the liver, and the serum TG content of the HFD group was significantly higher than that in the SD group, indicating the homeostasis imbalance in the lipid. We successfully obtained the fatty liver model we needed. However, the addition of Rosi and GW9 had no significant effect on the growth and metabolism of tilapia in low fat or high lipid background. Therefore, two experiments were carried out by intraperitoneal injection of this more direct method. The amount of Rosiglitazone injection was 0 (DMSO), 0.6mg/kg, 18mg/kg and 5. 4 mg/kg; the amount of GW9662 injection was 0 (DMSO), 0.4mg/kg, 12mg/kg, and 36 mg/kg, and the injection of GW9662 on serum triglyceride (Triglyceride, TG), free fatty acids (Free fatty acid), and glucose had no significant effect on blood glucose. TG content increased the content of FFA. Fluorescence quantitative PCR analysis found that fatty acid synthesis in the liver, TG synthesis and VLDL key protein gene expression decreased, indicating that the decrease of TG content was mainly attributed to the decrease of liver lipid synthesis and secretion, and the increase of FFA was attributed to the enhancement of lipid hydrolysis in the muscles. But Rosi did not affect the PPAR gamma in adipose tissue. The expression in the liver and muscle, which regulates the post transcriptional modification of PPAR gamma, affects its protein activity and causes the corresponding physiological effects. The results suggest that short term Rosi injection can promote the cloning of the lipid hydrolysis of the.4. PPAR gamma target gene ACOX1 of the tilapia muscle and its nutritional regulation of peroxisomes. The metabolic activity in peroxisomes is regulated by PPAR gamma, and the ester acyl coenzyme A oxidase 1 (Acayl-CoA Oxidase 1, ACOX1) is the first rate limiting enzyme of peroxisome beta oxidation, and it participates in the catalytic process of ester acyl coenzyme A to 2- CIS enyl coenzyme A, and peroxisome beta oxidation process as a letter. The synthesis of molecule.ACOX1 is one of the target genes of PPAR gamma. Therefore, it is important to elucidate the structure and function of ACOX1 for the study of how PPAR gamma regulates the synthesis of signal molecules of peroxidation objects. In this study, the ACOX1 gene was cloned in Nile tilapia for the first time. It was found to have two transcripts, named ACOXli1, respectively. And ACOXli2, their encoded proteins are composed of 661 amino acid residues. The two subtype of the encoding region is encoded by the exon E3b, which consists of 14 exons, and the ACOXli2, the ACOXli2, is encoded by E3a. The homology analysis shows that the conservatism of ACOXli1 is significantly higher than that of ACOXli2, suggesting that ACOXli1's function may be more than ACOXli2. The mRNA distribution pattern of the two groups showed that the expression of ACOXi1 in the liver was the highest, followed by the kidney and the brain, and the expression of ACOXli2 in the kidney was the highest, followed by the liver. The expression of the two subtypes in the white muscles, gills, brain and kidneys was significantly different, and the ACOXli2 was significantly higher in the white muscles and kidneys than in the ACOXlil, while in the brain, the brain was significantly higher than that in the brain. In contrast to the gills, in the 36 hours of hunger, the nutritional status of 1,3,8,24 hours after full food was found: the two subtypes made the same response pattern in the liver, the kidney ACOXlil and the liver response patterns converged, and the ACOXli2 was not affected by the nutritional status. The experimental results showed that the two subtypes of the ACOXl gene were in the ridge. Different functions may be exercised in vertebroid animals.
【学位授予单位】：华东师范大学
【学位级别】：博士
【学位授予年份】：2016
【分类号】：Q953
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本文编号：2174136
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