皱纹盘鲍（Haliotis discus hannai Ino.）L-古洛糖酸-1，4-内酯氧化酶（GLO）的研究
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摘要
本论文首先介绍了脊椎动物维生素C合成关键酶——L-古洛糖酸-1,4-内酯氧化酶(GLO)研究进展,包括脊椎动物中GLO的性质,影响脊椎动物GLO活性的因素,GLO在脊椎动物中的存在、分布以及与脊椎动物进化的关系。然后,通过分子生物学手段,围绕皱纹盘鲍(Haliotis discus hannai Ino.)是否具有GLO活性,是否具有VC合成能力这一主题进行了研究。主要研究内容包括：(1)皱纹盘鲍GLO基因的克隆和组织及不同发育阶段的表达；(2)饲料维生素C对皱纹盘鲍成鲍生长、各组织抗坏血酸含量以及GLO mRNA表达量的影响；(3)皱纹盘鲍VC缺乏条件下肝胰腺和肾脏差异表达cDNA文库的构建。研究结果总结如下：
(1)皱纹盘鲍GLO基因的克隆和组织及不同发育阶段的表达。配制VC0.0 mg/kg的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。以肾脏组织mRNA为模板,通过简并引物PCR法克隆GLO核心片段,通过RACE法调取基因全长。通过特异引物PCR和荧光实时定量PCR法确定皱纹盘鲍不同组织和不同发育阶段的表达。结果如下：获得了皱纹盘鲍GLO cDNA全长序列,包括1606个核苷酸,其中开放阅读框包括1365个核苷酸,编码一个由454个氨基酸组成的蛋白质,其氨基酸序列与脊椎动物的GLO序列有很高的同源性(>48%),确定其为皱纹盘鲍GLO基因；GLO mRNA在包括肝胰腺、肾脏、外套膜、肌肉、鳃和血细胞等6个组织中都有表达；GLO mRNA在从担轮幼虫到幼鲍的7个发育阶段都有表达,并且相对表达量在上足分化幼虫中达到最高,呈现随幼体发育先升高后下降的趋势。本研究在国际上首次从无脊椎动物中克隆到GLO全长序列,证明皱纹盘鲍具有VC合成能力。
(2)饲料维生素C对皱纹盘鲍成鲍生长、各组织抗坏血酸含量以及GLOmRNA表达量的影响。配制VC0.0,70.3,829.8和4967.5 mg/kg的共4个水平的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。结果表明,饲料中不同VC含量对皱纹盘鲍成鲍的存活率,增重率,贝壳日增长率等生长指标没有显著影响(p>0.05)。皱纹盘鲍不同组织间抗坏血酸含量存在极显著差异(p<0.01),鳃在各处理组均极显著的高于其他各组织(p<0.01)；而血清除在70.3mg/kg处理组显著低于外套膜(p<0.05)外,其他均极显著的低于其他组织(p<0.01)；总体而言,皱纹盘鲍组织抗坏血酸含量为鳃>肾脏、肝胰腺>外套膜、肌肉>血清。饲料中不同添加水平的维生素C对皱纹盘鲍各个组织抗坏血酸含量产生不同的影响,对肾脏、外套膜、鳃和血清抗坏血酸含量没有显著影响(p>0.05)；而对肌肉和肝胰腺抗坏血酸含量产生显著影响(p<0.05)：当饲料中维生素C含量达到4967.5 mg/kg时,肌肉中的抗坏血酸含量极显著的高于0.0 mg/kg处理组(p<0.01),显著高于829.8 mg/kg处理组(p<0.05)；肝胰腺中的抗坏血酸含量显著高于0.0 mg/kg处理组(p<0.05)。不同组织中两种看家基因β-actin和核糖体蛋白S9 mRNA的表达量均存在极显著差异(p<0.01),总体而言,β-actin mRNA的表达量血细胞>鳃、肌肉、外套膜、肾脏>肝胰腺；核糖体蛋白S9 mRNA的表达量鳃>血细胞、外套膜、肾脏、肝胰腺>肌肉。不同饲料维生素C对皱纹盘鲍β-actin mRNA的表达量在肝胰腺、肾脏、肌肉和外套膜中不受饲料维生素C含量的影响(p>0.05),但在鳃和血细胞中受饲料维生素C含量的影响(p<0.05)；核糖体蛋白S9 mRNA的表达量在肝胰腺、肾脏、肌肉、外套膜和鳃中均无显著影响(p>0.05)。以核糖体蛋白S9为内参基因时,饲料维生素C对各组织中GLOmRNA相对表达量均没有影响(p>0.05)；以β-actin为内参基因时,除鳃之外的其他各组织中GLO mRNA相对表达量均不受饲料维生素C的影响(p>0.05)；在鳃中,O.Omg/kg处理组显著高于829.8mg/kg处理组(p<0.05)。各组织间GLOmRNA相对表达量存在极显著差异(p<0.01),总体而言,GLO mRNA相对表达量肾脏>鳃>血细胞、外套膜、肝胰腺>肌肉。
(3)皱纹盘鲍VC缺乏条件下肝胰腺和肾脏差异表达cDNA文库的构建。配制了维生素C缺乏(0 mg/kg)和正常(4967.5 mg/kg)2个水平的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。采用抑制消减杂交法(suppression subtractive hybridization, SSH),构建了维生素C缺乏条件下肝胰腺和肾脏组织差异表达cDNA文库。消减杂交效率分析显示,构建的2个差异表达cDNA文库中差异表达的基因分别至少被富集了25和25-10倍。从文库中随机挑选阳性克隆测序。在肝胰腺差异表达cDNA文库中,获得了63个基因片段。在肾脏差异表达cDNA文库中,获得了39个基因片段。这些片段有可能为VC缺乏条件下差异表达的基因，这些片段的获得为进一步从分子水平研究皱纹盘鲍VC代谢机理奠定基础。
A detailed review of studying status on L-gulono-1,4-lactone oxidase (GLO), a key enzyme required for biosynthesis of ascorbic acid in vertebrate. A series of experiments were conducted to elucidate whether or not the abalone(Haliotis discus hannai Ino.) has GLO activity and the ability to synthesize VC by using molecular biology techniques. The current studies include the followings:(1) Isolation, cDNA cloning and gene expression of GLO in different tissue and development phase of abalone (Haliotis discus hannai Ino.). (2) The effects of dietary vitamin C on growth, tissue concentration of ascorbic acid and GLO mRNA relative expression level in different tissue of Haliotis discus hannai Ino..(3) Construction of the hepatopancreas and kidney cDNA subtractive library of Haliotis discus hannai Ino. fed with vitamin C-deficiency diet. The results are summarized as follows.
(1) The study was conducted to isolate and clone full-length cDNA sequence of GLO from Haliotis discus hannai Ino., and to investigate the relative expression level of GLO mRNA in different tissue and different development phase. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified vitamin C deficiency diet (VC 0.0 mg/kg). The full-length sequence (1606 bp) was determined from kidney of abalone fed with vitamin C deficiency diet. The open reading frames contained 454 amino acid which exhibited high amino acid identity (>48%) with vertebrate GLOs. The GLO mRNA expression was demonstrated in the abalone hepatopancreas, kidney, muscle, mantle, gill, blood cells and different development phase from trochosphere to 140d-old juvenile abalone. The GLO mRNA relative expression level of different development phase was also conducted by real-time quantitative PCR (RT-PCR). The result shows that the GLO mRNA relative expression level reaches the highest in the larva with pallial tentacle. W ith the development of abalone, relative expression level increased first and then decreased. The result suggested Haliotis discus hannai Ino. has the ability to synthesize VC.
(2) The study was conducted to investigate the effects of dietary vitamin C on growth, tissue ascorbic acid concentration and GLO mRNA relative expression level in different tissue of Haliotis discus hannai Ino.. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified diets containing 4 levels of vitamin C:0.0,70.3,829.8 and 4967.5 mg/kg. The result shows that survival and growth were not significantly affected by dietary treatments (p>0.05). Ascorbic acid concentration had significantly difference in different tissue (p<0.01). It was significantly higher (p<0.01) in gill and lower (p<0.05) in serum than the other tissues. Ascorbic acid concentration in different tissue ranked as follows:gill>kidney and hepatopancreas> muscle and mantle>serum. Ascorbic acid concentration was not significantly affected in kidney, gill, mantle and serum by dietary vitamin C (p>0.05), but significantly increased in hepatopancreas and muscle (p<0.05) with the increase dietary vitamin C. The mRNA relative expression level of two species of housekeeping geneβ-actin and ribosome protein S9 had significantly difference in different tissue(p<0.01). The mRNA relative expression level ofβ-actin in different tissue ranked as follows:blood cells> gill, muscle, mantle, and kidney> hepatopancreas. The mRNA relative expression level of ribosome protein S9 in different tissue ranked as follows:gill> blood cells, mantle, kidney and hepatopancreas> muscle. The mRNA relative expression level ofβ-actin was not significantly affected in hepatopancreas, kidney, mantle and muscle (p>0.05), but significantly affected in gill and blood cell (p<0.05) by dietary vitamin C. The mRNA relative expression level of ribosome protein S9 was not significantly affected in every tissue by dietary vitamin C (p>0.05). The mRNA relative expression level of GLO was not significantly affected in every tissue by dietary vitamin C when ribosome protein S9 was selected as reference gene (p>0.05). The mRNA relative expression level of GLO was not significantly affected in other tissues except gill by dietary vitamin C whenβ-actin was selected as reference gene (p>0.05). The mRNA relative expression level of GLO had significantly difference in different tissue(p<0.01).It was ranked as follows:kidney> gill, blood cells, mantle, and hepatopancreas> muscle.
(3) The study was conducted to clone the diffierentially expressed genes in the hepatopancreas and kidney of abalone (Haliotis discus hannai Ino.) under vitamin C deficiency. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified diets containing 2 levels of vitamin C:0.0 mg/kg and 4967.5 mg/kg. The cDNA subtractive library of the hepatopancreas·and kidney of abalone vitamin C deficiency-treated was constructed using the method of suppression subtractive hybridization Analysis of subtraction efficiency shows that diffierentially expressed genes had at least 25 and 25-10 - fold enrichment in two cDNA subtractive librarys respectively. Sixty three and thirty nine cDNA fragments which might be differentially expressed genes in the vitamin C deficiency group were obtained from the hepatopancreas and kidney cDNA subtractive library respectively by randomly picking and sequencing positive clones. This study plays an important role in researching the vitamin C metabolic mechanism of abalone at the molecular level.
(1)皱纹盘鲍GLO基因的克隆和组织及不同发育阶段的表达。配制VC0.0 mg/kg的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。以肾脏组织mRNA为模板,通过简并引物PCR法克隆GLO核心片段,通过RACE法调取基因全长。通过特异引物PCR和荧光实时定量PCR法确定皱纹盘鲍不同组织和不同发育阶段的表达。结果如下：获得了皱纹盘鲍GLO cDNA全长序列,包括1606个核苷酸,其中开放阅读框包括1365个核苷酸,编码一个由454个氨基酸组成的蛋白质,其氨基酸序列与脊椎动物的GLO序列有很高的同源性(>48%),确定其为皱纹盘鲍GLO基因；GLO mRNA在包括肝胰腺、肾脏、外套膜、肌肉、鳃和血细胞等6个组织中都有表达；GLO mRNA在从担轮幼虫到幼鲍的7个发育阶段都有表达,并且相对表达量在上足分化幼虫中达到最高,呈现随幼体发育先升高后下降的趋势。本研究在国际上首次从无脊椎动物中克隆到GLO全长序列,证明皱纹盘鲍具有VC合成能力。
(2)饲料维生素C对皱纹盘鲍成鲍生长、各组织抗坏血酸含量以及GLOmRNA表达量的影响。配制VC0.0,70.3,829.8和4967.5 mg/kg的共4个水平的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。结果表明,饲料中不同VC含量对皱纹盘鲍成鲍的存活率,增重率,贝壳日增长率等生长指标没有显著影响(p>0.05)。皱纹盘鲍不同组织间抗坏血酸含量存在极显著差异(p<0.01),鳃在各处理组均极显著的高于其他各组织(p<0.01)；而血清除在70.3mg/kg处理组显著低于外套膜(p<0.05)外,其他均极显著的低于其他组织(p<0.01)；总体而言,皱纹盘鲍组织抗坏血酸含量为鳃>肾脏、肝胰腺>外套膜、肌肉>血清。饲料中不同添加水平的维生素C对皱纹盘鲍各个组织抗坏血酸含量产生不同的影响,对肾脏、外套膜、鳃和血清抗坏血酸含量没有显著影响(p>0.05)；而对肌肉和肝胰腺抗坏血酸含量产生显著影响(p<0.05)：当饲料中维生素C含量达到4967.5 mg/kg时,肌肉中的抗坏血酸含量极显著的高于0.0 mg/kg处理组(p<0.01),显著高于829.8 mg/kg处理组(p<0.05)；肝胰腺中的抗坏血酸含量显著高于0.0 mg/kg处理组(p<0.05)。不同组织中两种看家基因β-actin和核糖体蛋白S9 mRNA的表达量均存在极显著差异(p<0.01),总体而言,β-actin mRNA的表达量血细胞>鳃、肌肉、外套膜、肾脏>肝胰腺；核糖体蛋白S9 mRNA的表达量鳃>血细胞、外套膜、肾脏、肝胰腺>肌肉。不同饲料维生素C对皱纹盘鲍β-actin mRNA的表达量在肝胰腺、肾脏、肌肉和外套膜中不受饲料维生素C含量的影响(p>0.05),但在鳃和血细胞中受饲料维生素C含量的影响(p<0.05)；核糖体蛋白S9 mRNA的表达量在肝胰腺、肾脏、肌肉、外套膜和鳃中均无显著影响(p>0.05)。以核糖体蛋白S9为内参基因时,饲料维生素C对各组织中GLOmRNA相对表达量均没有影响(p>0.05)；以β-actin为内参基因时,除鳃之外的其他各组织中GLO mRNA相对表达量均不受饲料维生素C的影响(p>0.05)；在鳃中,O.Omg/kg处理组显著高于829.8mg/kg处理组(p<0.05)。各组织间GLOmRNA相对表达量存在极显著差异(p<0.01),总体而言,GLO mRNA相对表达量肾脏>鳃>血细胞、外套膜、肝胰腺>肌肉。
(3)皱纹盘鲍VC缺乏条件下肝胰腺和肾脏差异表达cDNA文库的构建。配制了维生素C缺乏(0 mg/kg)和正常(4967.5 mg/kg)2个水平的精制饲料,在室内流水养殖系统中,投喂初重为74.32±0.43 g,初始壳长为84.36±0.24 mm的皱纹盘鲍成鲍,养殖周期170天。采用抑制消减杂交法(suppression subtractive hybridization, SSH),构建了维生素C缺乏条件下肝胰腺和肾脏组织差异表达cDNA文库。消减杂交效率分析显示,构建的2个差异表达cDNA文库中差异表达的基因分别至少被富集了25和25-10倍。从文库中随机挑选阳性克隆测序。在肝胰腺差异表达cDNA文库中,获得了63个基因片段。在肾脏差异表达cDNA文库中,获得了39个基因片段。这些片段有可能为VC缺乏条件下差异表达的基因，这些片段的获得为进一步从分子水平研究皱纹盘鲍VC代谢机理奠定基础。
A detailed review of studying status on L-gulono-1,4-lactone oxidase (GLO), a key enzyme required for biosynthesis of ascorbic acid in vertebrate. A series of experiments were conducted to elucidate whether or not the abalone(Haliotis discus hannai Ino.) has GLO activity and the ability to synthesize VC by using molecular biology techniques. The current studies include the followings:(1) Isolation, cDNA cloning and gene expression of GLO in different tissue and development phase of abalone (Haliotis discus hannai Ino.). (2) The effects of dietary vitamin C on growth, tissue concentration of ascorbic acid and GLO mRNA relative expression level in different tissue of Haliotis discus hannai Ino..(3) Construction of the hepatopancreas and kidney cDNA subtractive library of Haliotis discus hannai Ino. fed with vitamin C-deficiency diet. The results are summarized as follows.
(1) The study was conducted to isolate and clone full-length cDNA sequence of GLO from Haliotis discus hannai Ino., and to investigate the relative expression level of GLO mRNA in different tissue and different development phase. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified vitamin C deficiency diet (VC 0.0 mg/kg). The full-length sequence (1606 bp) was determined from kidney of abalone fed with vitamin C deficiency diet. The open reading frames contained 454 amino acid which exhibited high amino acid identity (>48%) with vertebrate GLOs. The GLO mRNA expression was demonstrated in the abalone hepatopancreas, kidney, muscle, mantle, gill, blood cells and different development phase from trochosphere to 140d-old juvenile abalone. The GLO mRNA relative expression level of different development phase was also conducted by real-time quantitative PCR (RT-PCR). The result shows that the GLO mRNA relative expression level reaches the highest in the larva with pallial tentacle. W ith the development of abalone, relative expression level increased first and then decreased. The result suggested Haliotis discus hannai Ino. has the ability to synthesize VC.
(2) The study was conducted to investigate the effects of dietary vitamin C on growth, tissue ascorbic acid concentration and GLO mRNA relative expression level in different tissue of Haliotis discus hannai Ino.. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified diets containing 4 levels of vitamin C:0.0,70.3,829.8 and 4967.5 mg/kg. The result shows that survival and growth were not significantly affected by dietary treatments (p>0.05). Ascorbic acid concentration had significantly difference in different tissue (p<0.01). It was significantly higher (p<0.01) in gill and lower (p<0.05) in serum than the other tissues. Ascorbic acid concentration in different tissue ranked as follows:gill>kidney and hepatopancreas> muscle and mantle>serum. Ascorbic acid concentration was not significantly affected in kidney, gill, mantle and serum by dietary vitamin C (p>0.05), but significantly increased in hepatopancreas and muscle (p<0.05) with the increase dietary vitamin C. The mRNA relative expression level of two species of housekeeping geneβ-actin and ribosome protein S9 had significantly difference in different tissue(p<0.01). The mRNA relative expression level ofβ-actin in different tissue ranked as follows:blood cells> gill, muscle, mantle, and kidney> hepatopancreas. The mRNA relative expression level of ribosome protein S9 in different tissue ranked as follows:gill> blood cells, mantle, kidney and hepatopancreas> muscle. The mRNA relative expression level ofβ-actin was not significantly affected in hepatopancreas, kidney, mantle and muscle (p>0.05), but significantly affected in gill and blood cell (p<0.05) by dietary vitamin C. The mRNA relative expression level of ribosome protein S9 was not significantly affected in every tissue by dietary vitamin C (p>0.05). The mRNA relative expression level of GLO was not significantly affected in every tissue by dietary vitamin C when ribosome protein S9 was selected as reference gene (p>0.05). The mRNA relative expression level of GLO was not significantly affected in other tissues except gill by dietary vitamin C whenβ-actin was selected as reference gene (p>0.05). The mRNA relative expression level of GLO had significantly difference in different tissue(p<0.01).It was ranked as follows:kidney> gill, blood cells, mantle, and hepatopancreas> muscle.
(3) The study was conducted to clone the diffierentially expressed genes in the hepatopancreas and kidney of abalone (Haliotis discus hannai Ino.) under vitamin C deficiency. Abalone, (initial weight:74.66±1.0 g; initial shell length:84.36±1.07 mm) were fed 170 days with purified diets containing 2 levels of vitamin C:0.0 mg/kg and 4967.5 mg/kg. The cDNA subtractive library of the hepatopancreas·and kidney of abalone vitamin C deficiency-treated was constructed using the method of suppression subtractive hybridization Analysis of subtraction efficiency shows that diffierentially expressed genes had at least 25 and 25-10 - fold enrichment in two cDNA subtractive librarys respectively. Sixty three and thirty nine cDNA fragments which might be differentially expressed genes in the vitamin C deficiency group were obtained from the hepatopancreas and kidney cDNA subtractive library respectively by randomly picking and sequencing positive clones. This study plays an important role in researching the vitamin C metabolic mechanism of abalone at the molecular level.
引文
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[1]Nishikimi M, Yagi K. Molecular basis for the deficiency in humans of gulonolactone oxidase, a key enzyme for ascorbic acid biosynthesis. American Journal of Clinical Nutrition.1991,54(6):1203-1208
[2]Fracalossi D M, AllenM E, Yuyanma L K, et al. Ascorbic acid biosynthesis in Amazonian Fish. Aquaculture,2001,192:321-332
[3]Moreau, R., Dabrowski, K. Body pool and synthesis of ascorbic acid in sea lamprey (Petromyzon marinus):an agnathan fish with gulonolactone oxidase activity. Proc. Natl. Acad. Sci.1998a.U.S.A.9510279-10282.
[4]Dabrowski K. Gulonolactone oxidase is missing in teleost fish — the direct spectrophotometric assay. Biological Chemistry Hoppe-Seyler,1990,371:207-214.
[5]Dabrowski K. Primitive Actinopterigian fishes can synthesize ascorbic acid. Experientia, 1994,50:745-748.
[6]Touhata K, Toyohara H, Mitani T et al. Distribution of L-gulono-1,4-lactone oxidase among fishes. Fish Science,1995,61:729-730.
[7]Maeland A, Waagbo R. Examination of the qualitative ability of some cold water marine teleosts to synthesise ascorbic acid. Comparative Biochemistry and Physiology-Part A: Molecular & Integrative Physiology,1998,121:249-255.
[8]Moreau R, Dabrowski K. Biosynthesis of ascorbic acid by extant actinopterygians. Journal of Fish Biology,2000,57:733-745
[9]Moreau, R., Kaushik, S.J., Dabrowski, K.,1996. Ascorbic acid status as affected by dietary treatment in the Siberian sturgeon Acipenser baeri Brandt:tissue concentration, mobilisation and L-gulonolactone oxidase activity. Fish Physiol. Biochem.15:431-438.
[10]Ikeda S, Sato M. Biochemcal studies on L-ascorbic acid in aquatic animalsⅢ. Biosybthesis of L-ascorbic acid by carp [J]. Bulletin of the Japanese Society of Scientific Fisheries,1964, 30:365-374.
[11]Sato M, Yoshinaka R, Yamamoto Y et al. Nonessentiality of ascorbic acid in the diet of carp. Bulletin of the Japanese Society of Scientific Fisheries,1978,44:1151-1156.
[12]Yamamoto Y, Sato M, Ikeda S. Existence of L-gulonolactone oxidase in some teleosts. Bulletin of the Japanese Society of Scientific Fisheries,1978,44:775-779.
[13]Soliman A K, Jauncey K, Roberts R J. Qualitative and quantitative identification of L-gulonolactone oxidase activity in some teleosts. Aquaculture and Fisheries Management, 1985,1:249-256.
[14]Thomas P, Bally M B, Neff J M. Influence of some environmental variables on the ascorbic acid status of mullet, Mugil cephalus L., tissues:Ⅱ. Seasonal fluctuations and biosynthetic ability. Journal of Fish Biology,1985,27:47-57.
[15]Chatterjee I B, Majumder A K, Nand B K, et al. Evolution and the Biosynthesis of Ascorbic Acid. Science,1973,182:1271-1272.
[16]Roy R N, Guha B C. Species Difference in regard to the Biosynthesis of Ascorbic Acid. Nature,1958,182:319-320.
[17]Chaudhuri C R, Chatterjee I B. L-ascorbic acid synthesis in birds:phylogenetic trend. Science,1969,164(878):435-436.
[18]Jenness, R., Birney, E.C., Ayaz, K.L.,1980. Variation of L-gulonolactone oxidase activity in placental mammals. Comp. Biochem. Physiol., Part B 67,195-204.
[19]Bimey E C, Jenness J J, Humer I D. Ascorbic acid biosynthesis in the mammalian kidney. Experientia,1979,35:1425-1426
[20]Chatterjee I B, Majumder A K, Nandi B K, et al. Synthesis and some major functions of vitamin C in animals. Annals of the New York Academy of Sciences.1975,258:24-47
[21]Burns J J. Missing Step in Man, Monkey and guinea pig required for the biosynthesis of L-ascorbic acid. Nature,1957,180:553.
[22]Koshizaka T, Nishikimi M, Ozawa T et al. Isolation and sequence analysis of a complementary DNA encoding rat liver L-gulono-gamma-lactone oxidase, a key enzyme for L-ascorbic acid biosynthesis. Journal of Biological Chemistry,1988,263 (4):1619-1621.
[23]Ha M N, Graham F L, D'Souza C K, et al. Functional rescue of vitamin C synthesis deficiency in human cells using adenoviral-based expression of murine L-gulono-γ-lactone oxidase. Genomics,2004,83:482-492.
[24]Nam Y K, Cho Y S, Douglas S E et al. Isolation and transient expression of a cDNA encoding l-gulono-γ-lactone oxidase, a key enzyme for l-ascorbic acid biosynthesis, from the tiger shark Scyliorhinus torazame. Aquaculture,2002,209:271-284
[25]Cho Y S, Douglas S E, Gallant J W, et al. Isolation and characterization of cDNA sequences of L-gulono-gamma-lactone oxidase, a key enzyme for biosynthesis of ascorbic acid, from extant primitive fish groups. Biochemical and Biophysical Research Communications,2007,147(2):178-190
[26]Mai K. Comparative studies on the nutrition of two species of abalone, Haliotis tuberculata L. and Haliotis discus hannai Ino.:VII. Effects of dietary vitamin C on survival, growth and tissue concentration of ascorbic acid. Aquaculture,1998,161:383-392
[27]Tan B., Mai K.. Zinc methionine and zinc sulfate as sources of dietary zinc for juvenile abalone, Haliotis discus hannai Ino.. Aquaculture,192,2001:67-84
[28]Uki, N., A. Kemuyama. T. Watanabe. Development of semipurified test diets for abalone. Bull. Jap. Soc. Sci.Fish.,1985,51:1825-1833.
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[31]周歧存,麦康森,谭北平,徐玮。维生素E对皱纹盘鲍幼鲍生长、存活及体成分的影响。海洋与湖沼,32(2),2001：125-131.
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[40]Nishikimi M, Yagi K. Molecular basis for the deficiency in humans of gulonolactone oxidase, a key enzyme for ascorbic acid biosynthesis. American Journal of Clinical Nutrition.1991,54(6):1203-1208
[41]Kiuchi K, Nishikimi M, Yagi K. Purification and characterization of L-gulonolactone oxidase from chicken kidney microsomes. Biochemistry,1982,21:5076-5082.
[42]Banhegyi G, Csala M, Braun L, Garzo T & Mandl J. Ascorbate synthesis-dependent glutathione consumption in mouse liver. FEBS Lett,1996,381:39-41.
[43]Halliwell, B., and Gutteridge, J. M. C.. "Free Radicals in Biology and Medicine."Clarendon Press, Oxford.1989.
[44]Arrigoni 0, De Tullio M C. Ascorbic acid:much more than just an antioxidant. Biochimica et Biophysica Acta,2002,1569(1-3):1-9.
[45]Moreau R, Dabrowski K. Alpha-tocopherol downregulates gulonolactone oxidase activity in sturgeon. Free Radical Biology and Medicine.2003,34(10):1326-32.
[46]Jenness R, Birney E C, Ayaz K L. Ascorbic acid and L-gulonolactone oxidase in lagomorphs. Comparative Biochemistry and Physiology,1978,61(3):395-399.
[47]R. Jenness, E.C. Bimey, K.L. Ayaz, D.M. Buzzell, Ontogenetic development of L-gulonolactoneoxidase activity in several vertebrates, Comp. Biochem. Physiol. B 78 1984 167-173.
[48]Nandi A, Mukhopadhyay C K, Ghosh M K et al. Evolutionary Significance of Vitamin C Biosynthesis in Terrestrial Vertebrates. Free Radical Biology and Medicine,1997, 22:1047-1054
[1]Nishikimi M, Yagi K. Molecular basis for the deficiency in humans of gulonolactone oxidase, a key enzyme for ascorbic acid biosynthesis. American Journal of Clinical Nutrition.1991, 54(6):1203-1208
[2]Jenness R, Birney E C, Ayaz K L. Ascorbic acid and L-gulonolactone oxidase in lagomorphs. Comparative Biochemistry and Physiology,1978,61(3):395-399.
[3]Tsao C S, Young M. Effect of exogenous ascorbic acid intake on biosynthesis of ascorbic acid in mice. Life Science,1989,45:1553-1557.
[4]Hooper C L, Maurice D V, Lightsey S F, et al. Factors affecting ascorbic acid (AsA) biosynthesis in chickens.Ⅱ. Effect of dietary AsA and strain of chicken. Journal of Animal Physiology and Animal Nutrition,2002,86:326-332
[5]Moreau R, Dabrowski K, Sato P H. Renal L-gulono-1,4-lactone oxidase activity as affected by dietary ascorbic acid in lake sturgeon (Acipenser fulvescens). Aquaculture,1999, 180:359-372
[6]Xie Z G, Niu C J, Zhang Z B et al. Dietary ascorbic acid may be necessary for enhancing the immune response in Siberian sturgeon (Acipenser baerii), a species capable of ascorbic acid biosynthesis. Comparative Biochemistry and Physiology-part A:Molecular & Integrative Physiology 2006,145:152-157
[7]Chatterjee I B, Majumder A K, Nand B K, et al. Evolution and the Biosynthesis of Ascorbic Acid. Science,1973,182:1271-1272.
[8]Mai K. Comparative studies on the nutrition of two species of abalone, Haliotis tuberculata L. and Haliotis discus hannai Ino.:VII. Effects of dietary vitamin C on survival, growth and tissue concentration of ascorbic acid. Aquaculture,1998,161:383-392
[9]Tan B., Mai K.. Zinc methionine and zinc sulfate as sources of dietary zinc for juvenile abalone, Haliotis discus hannai Ino.. Aquaculture,192,2001:67-84
[10]Uki, N., A. Kemuyama. T. Watanabe. Development of semipurified test diets for abalone. Bull. Jap. Soc. Sci.Fish.,1985,51:1825-1833.
[11]Mai, K., J. P. Mercer, J. Donlon. Comparative studies on the nutrition of two species of abalone,Haliotis mberculam L and Haliotis discus hannai Ino. IV. Optimum dietary protein level for growth. Aquaculture,1995a,136:165-180.
[12]Mai, K., J. P. Mercer, J. Donlon. Comparative studies on the nutrition of two species of abalone,Haliotis tuberculata L and Haliotis discus hannai Ino.Ⅱ. Response of abalone to various levels of dietary lipid. Aquaculture,1995b,134:65-80.
[13]周歧存,麦康森,谭北平,徐玮。维生素E对皱纹盘鲍幼鲍生长、存活及体成分的影响。海洋与湖沼,32(2),2001：125-131.
[14]Xiaojie Wang, Kang-Woong Kim, Sungchul C. Bai. Comparison of L-ascorbyl-2-monophosphate-Ca with L-ascorbyl-2-monophosphate-Na/Ca on growth and tissue ascorbic acid concentrations in Korean rockfish (Sebastes schlegeli). Aquaculture 225 (2003) 387-395.
[15]Qinghui Ai, Kangsen Mai, Chunxiao Zhang, Wei Xu, Qingyuan Duan, Beiping Tan, Zhiguo Liufu. Effects of dietary vitamin C on growth and immune response of Japanese seabass, Lateolabrax japonicus. Aquaculture 242 (2004) 489-500.
[16]Fracalossi D M, AllenM E, Yuyanma L K, et al. Ascorbic acid biosynthesis in Amazonian Fish. Aquaculture,2001,192:321-332
[17]Moreau, R., Dabrowski, K. Body pool and synthesis of ascorbic acid in sea lamprey (Petromyzon marinus):an agnathan fish with gulonolactone oxidase activity. Proc. Natl. Acad. Sci.1998a.U.S.A.9510279-10282.
[18]Dabrowski K. Gulonolactone oxidase is missing in teleost fish — the direct spectrophotometric assay. Biological Chemistry Hoppe-Seyler,1990,371:207-214.
[19]Dabrowski K. Primitive Actinopterigian fishes can synthesize ascorbic acid. Experientia, 1994,50:745-748.
[20]Touhata K, Toyohara H, Mitani T et al. Distribution of L-gulono-1,4-lactone oxidase among fishes. Fish Science,1995,61:729-730.
[21]Maeland A, Waagbo R. Examination of the qualitative ability of some cold water marine teleosts to synthesise ascorbic acid. Comparative Biochemistry and Physiology-Part A: Molecular & Integrative Physiology,1998,121:249-255.
[22]Moreau R, Dabrowski K. Biosynthesis of ascorbic acid by extant actinopterygians. Journal of Fish Biology,2000,57:733-745
[23]Moreau, R., Kaushik, S.J., Dabrowski, K.,1996. Ascorbic acid status as affected by dietary treatment in the Siberian sturgeon Acipenser baeri Brandt:tissue concentration, mobilisation and L-gulonolactone oxidase activity. Fish Physiol. Biochem.15:431-438.
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[6]Touhata K, Toyohara H, Mitani T et al. Distribution of L-gulono-1,4-lactone oxidase among fishes. Fish Science,1995,61:729-730.
[7]Maeland A, Waagbo R. Examination of the qualitative ability of some cold water marine teleosts to synthesise ascorbic acid. Comparative Biochemistry and Physiology-Part A: Molecular & Integrative Physiology,1998,121:249-255.
[8]Moreau R, Dabrowski K. Biosynthesis of ascorbic acid by extant actinopterygians. Journal of Fish Biology,2000,57:733-745
[9]Moreau, R., Kaushik, S.J., Dabrowski, K.,1996. Ascorbic acid status as affected by dietary treatment in the Siberian sturgeon Acipenser baeri Brandt:tissue concentration, mobilisation and L-gulonolactone oxidase activity. Fish Physiol. Biochem.15:431-438.
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[11]Sato M, Yoshinaka R, Yamamoto Y et al. Nonessentiality of ascorbic acid in the diet of carp. Bulletin of the Japanese Society of Scientific Fisheries,1978,44:1151-1156.
[12]Yamamoto Y, Sato M, Ikeda S. Existence of L-gulonolactone oxidase in some teleosts. Bulletin of the Japanese Society of Scientific Fisheries,1978,44:775-779.
[13]Soliman A K, Jauncey K, Roberts R J. Qualitative and quantitative identification of L-gulonolactone oxidase activity in some teleosts. Aquaculture and Fisheries Management, 1985,1:249-256.
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[17]Chaudhuri C R, Chatterjee I B. L-ascorbic acid synthesis in birds:phylogenetic trend. Science,1969,164(878):435-436.
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[21]Burns J J. Missing Step in Man, Monkey and guinea pig required for the biosynthesis of L-ascorbic acid. Nature,1957,180:553.
[22]Koshizaka T, Nishikimi M, Ozawa T et al. Isolation and sequence analysis of a complementary DNA encoding rat liver L-gulono-gamma-lactone oxidase, a key enzyme for L-ascorbic acid biosynthesis. Journal of Biological Chemistry,1988,263 (4):1619-1621.
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[1]Nishikimi M, Yagi K. Molecular basis for the deficiency in humans of gulonolactone oxidase, a key enzyme for ascorbic acid biosynthesis. American Journal of Clinical Nutrition.1991, 54(6):1203-1208
[2]Jenness R, Birney E C, Ayaz K L. Ascorbic acid and L-gulonolactone oxidase in lagomorphs. Comparative Biochemistry and Physiology,1978,61(3):395-399.
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[4]Hooper C L, Maurice D V, Lightsey S F, et al. Factors affecting ascorbic acid (AsA) biosynthesis in chickens.Ⅱ. Effect of dietary AsA and strain of chicken. Journal of Animal Physiology and Animal Nutrition,2002,86:326-332
[5]Moreau R, Dabrowski K, Sato P H. Renal L-gulono-1,4-lactone oxidase activity as affected by dietary ascorbic acid in lake sturgeon (Acipenser fulvescens). Aquaculture,1999, 180:359-372
[6]Xie Z G, Niu C J, Zhang Z B et al. Dietary ascorbic acid may be necessary for enhancing the immune response in Siberian sturgeon (Acipenser baerii), a species capable of ascorbic acid biosynthesis. Comparative Biochemistry and Physiology-part A:Molecular & Integrative Physiology 2006,145:152-157
[7]Chatterjee I B, Majumder A K, Nand B K, et al. Evolution and the Biosynthesis of Ascorbic Acid. Science,1973,182:1271-1272.
[8]Mai K. Comparative studies on the nutrition of two species of abalone, Haliotis tuberculata L. and Haliotis discus hannai Ino.:VII. Effects of dietary vitamin C on survival, growth and tissue concentration of ascorbic acid. Aquaculture,1998,161:383-392
[9]Tan B., Mai K.. Zinc methionine and zinc sulfate as sources of dietary zinc for juvenile abalone, Haliotis discus hannai Ino.. Aquaculture,192,2001:67-84
[10]Uki, N., A. Kemuyama. T. Watanabe. Development of semipurified test diets for abalone. Bull. Jap. Soc. Sci.Fish.,1985,51:1825-1833.
[11]Mai, K., J. P. Mercer, J. Donlon. Comparative studies on the nutrition of two species of abalone,Haliotis mberculam L and Haliotis discus hannai Ino. IV. Optimum dietary protein level for growth. Aquaculture,1995a,136:165-180.
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[13]周歧存,麦康森,谭北平,徐玮。维生素E对皱纹盘鲍幼鲍生长、存活及体成分的影响。海洋与湖沼,32(2),2001：125-131.
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[19]Dabrowski K. Primitive Actinopterigian fishes can synthesize ascorbic acid. Experientia, 1994,50:745-748.
[20]Touhata K, Toyohara H, Mitani T et al. Distribution of L-gulono-1,4-lactone oxidase among fishes. Fish Science,1995,61:729-730.
[21]Maeland A, Waagbo R. Examination of the qualitative ability of some cold water marine teleosts to synthesise ascorbic acid. Comparative Biochemistry and Physiology-Part A: Molecular & Integrative Physiology,1998,121:249-255.
[22]Moreau R, Dabrowski K. Biosynthesis of ascorbic acid by extant actinopterygians. Journal of Fish Biology,2000,57:733-745
[23]Moreau, R., Kaushik, S.J., Dabrowski, K.,1996. Ascorbic acid status as affected by dietary treatment in the Siberian sturgeon Acipenser baeri Brandt:tissue concentration, mobilisation and L-gulonolactone oxidase activity. Fish Physiol. Biochem.15:431-438.
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[27]Soliman A K, Jauncey K, Roberts R J. Qualitative and quantitative identification of L-gulonolactone oxidase activity in some teleosts. Aquaculture and Fisheries Management, 1985,1:249-256.
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[29]Chaudhuri C R, Chatterjee I B. L-ascorbic acid synthesis in birds:phylogenetic trend. Science,1969,164(878):435-436.
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