大黄鱼生长激素基因的分析及其在大肠杆菌中的表达
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摘要
为了解决目前大黄鱼（Pseudosciaena crocea，pc）等名贵经济鱼类在养殖
中存在的养殖周期长，饵料利用率低，养殖生产成本较高的问题，急需开发鱼类
本身的基因产物以改良鱼类养殖饲料。鱼类生长激素（growth hormone，GH）是
鱼类脑垂体细胞分泌的多肽激素，具有增强食欲、提高饲料转化率的作用。天然
鱼 GH 含量极低，分离纯化成本较高。基因工程技术为生产大量低成本的鱼生长
激素创造了条件，且实验证明重组鱼 GH 具有天然鱼 GH 的功能，却不会对人体产
生激素副作用。因此，生产基因重组大黄鱼 GH 在养殖领域具有重要的应用价值。
本论文分析了最新克隆的 pcGH 基因的特征，成功实现了 pcGH 在大肠杆菌中
的胞内表达和周质表达，并进一步探讨了影响 pcGH 在大肠杆菌中表达水平的各
种因素。
通过与数据库中同类蛋白序列的比较，发现最新克隆的 pcGH 分别在不同位
置与数据库中的 pcGH 及黑斑红鲈（Sciaenops ocellata）GH 有高度同源性，推
断这个 pcGH 基因来源于多个近源鱼种的杂交体。这一发现将有助于进一步研究
分析鱼类 GH 的活性部位及其突变对鱼类生长的影响。
通过 PCR 引物设计，在 pcGH 基因两端引入两个酶切位点，将其插入到大肠
杆菌表达载体 pET-22b(+)中，构建胞内表达质粒 pG2。将 pG2 转化大肠杆菌
BL21(DE3)和 Rosetta(DE3)。SDS-PAGE 分析发现，pG2/Rosetta(DE3)在诱导后对
应于 pcGH 分子量的位置有特异蛋白带随时间增浓，但在 pG2/BL21(DE3)中该位
置蛋白带浓度变化不明显。说明了 Rosetta(DE3)菌株有助于改善 pcGH 中大肠杆
菌稀有密码子对表达水平的影响。pcGH2/Roseta(DE3)的表达产物为可溶性蛋白；
该产物用硫酸铵分级沉降和阳离子交换层析分离纯化，达到电泳纯。
通过酶切位点的选择，使 pcGH 基因插入 pET-22b(+)载体上 pel B 分泌肽编
码序列的下游，构建成功周质表达质粒 pG3；将 pG3 转化大肠杆菌 Rosetta(DE3)，
诱导表达后分离周质蛋白，发现部分重组蛋白在周质中表达。
本文还从转录水平和翻译水平系统、完整地探讨了影响 pcGH 在大肠杆菌中
表达水平的各种可能因素，分析了进一步提高重组大黄鱼 GH 的产量的可能方法。
这些工作对重组 pcGH 的生产和应用具有指导意义。
Problems such as low growth rate and low conversion rate of feedstuff exist in
the culturing of Pseudosciaena crocea and other economical marine fishes. Therefore,
it is economically important to exploit some genetic products of fish itself to improve
the properties of feedstuff. Fish growth hormone (fGH) is a polypeptide produced by
the pituitary cells of teleosts. It has the capacity to enhance the appetite and feed
conversion efficiency. However, the content of native fGH in pituitary is extremely
low and the expenditure of purification is too high. Fortunately, it is possible to
produce recombinant fGHs abundantly. And they have been proved in many cases the
same function as the native ones without any side effect to human. Therefore, it is
valuable to investigate the production of recombinant Pseudosciaena crocea GH
(pcGH).
In this research, the sequence of a newly cloned pcGH gene was analysed. Then
recombinant pcGH was produced both in intracellular and periplasmic manner. And
the factors effecting the expression level of the Escherichia coli were discussed.
The newly cloned pcGH showed high homogeneity to pcGH and Sciaenops
ocellata GH from GenBank in different regions respectively. Therefore this gene was
presumed to be the hybrid of several species of fish. This supposition will contribute
to find out the conserved regions of fGH. It'll also help to investigate the influence of
genetic mutation of GH on the growth of Pseudosciaena crocea.
The pcGH gene was cloned into pET-22b(+) for intracellular expression.
Recombinant plasmid (pG2) were transformed into E. coli BL21(DE3) and
Rosetta(DE3). SDS-PAGE analysis showed that a specific protein band with the same
molecular weight as theoretical one of pcGH was induced in pG2/Rosetta(DE3); but
in the case of pG2/BL21(DE3), this band was difficult to be found. Therefore, it’s
concluded that Rosetta(DE3) can help to lighten the effect of rare codon on the
expression level in E. coli. The product of pG2/Rosetta(DE3) was proved soluble,
then it was purified to be electrophoretically pure by ammonium sulfate precipitation
and cation exchange chromatography.
Another pair of primer was designed to insert the gene of pcGH into the
II
downstream of pel B signal peptide in pET-22b(+). The recombinant plasmid was
transformed into E. coli Rosetta(DE3). With IPTG induction, part of the recombinant
pGH was secreted into the periplasm of transformants.
Factors effecting the expression level of pcGH in E. coli, both on transcription
level and translation level, were discussed integrallty and systemically. Approches to
improve the expression level were put forward .
These works may have potential value in the production and application of
pcGH.
中存在的养殖周期长，饵料利用率低，养殖生产成本较高的问题，急需开发鱼类
本身的基因产物以改良鱼类养殖饲料。鱼类生长激素（growth hormone，GH）是
鱼类脑垂体细胞分泌的多肽激素，具有增强食欲、提高饲料转化率的作用。天然
鱼 GH 含量极低，分离纯化成本较高。基因工程技术为生产大量低成本的鱼生长
激素创造了条件，且实验证明重组鱼 GH 具有天然鱼 GH 的功能，却不会对人体产
生激素副作用。因此，生产基因重组大黄鱼 GH 在养殖领域具有重要的应用价值。
本论文分析了最新克隆的 pcGH 基因的特征，成功实现了 pcGH 在大肠杆菌中
的胞内表达和周质表达，并进一步探讨了影响 pcGH 在大肠杆菌中表达水平的各
种因素。
通过与数据库中同类蛋白序列的比较，发现最新克隆的 pcGH 分别在不同位
置与数据库中的 pcGH 及黑斑红鲈（Sciaenops ocellata）GH 有高度同源性，推
断这个 pcGH 基因来源于多个近源鱼种的杂交体。这一发现将有助于进一步研究
分析鱼类 GH 的活性部位及其突变对鱼类生长的影响。
通过 PCR 引物设计，在 pcGH 基因两端引入两个酶切位点，将其插入到大肠
杆菌表达载体 pET-22b(+)中，构建胞内表达质粒 pG2。将 pG2 转化大肠杆菌
BL21(DE3)和 Rosetta(DE3)。SDS-PAGE 分析发现，pG2/Rosetta(DE3)在诱导后对
应于 pcGH 分子量的位置有特异蛋白带随时间增浓，但在 pG2/BL21(DE3)中该位
置蛋白带浓度变化不明显。说明了 Rosetta(DE3)菌株有助于改善 pcGH 中大肠杆
菌稀有密码子对表达水平的影响。pcGH2/Roseta(DE3)的表达产物为可溶性蛋白；
该产物用硫酸铵分级沉降和阳离子交换层析分离纯化，达到电泳纯。
通过酶切位点的选择，使 pcGH 基因插入 pET-22b(+)载体上 pel B 分泌肽编
码序列的下游，构建成功周质表达质粒 pG3；将 pG3 转化大肠杆菌 Rosetta(DE3)，
诱导表达后分离周质蛋白，发现部分重组蛋白在周质中表达。
本文还从转录水平和翻译水平系统、完整地探讨了影响 pcGH 在大肠杆菌中
表达水平的各种可能因素，分析了进一步提高重组大黄鱼 GH 的产量的可能方法。
这些工作对重组 pcGH 的生产和应用具有指导意义。
Problems such as low growth rate and low conversion rate of feedstuff exist in
the culturing of Pseudosciaena crocea and other economical marine fishes. Therefore,
it is economically important to exploit some genetic products of fish itself to improve
the properties of feedstuff. Fish growth hormone (fGH) is a polypeptide produced by
the pituitary cells of teleosts. It has the capacity to enhance the appetite and feed
conversion efficiency. However, the content of native fGH in pituitary is extremely
low and the expenditure of purification is too high. Fortunately, it is possible to
produce recombinant fGHs abundantly. And they have been proved in many cases the
same function as the native ones without any side effect to human. Therefore, it is
valuable to investigate the production of recombinant Pseudosciaena crocea GH
(pcGH).
In this research, the sequence of a newly cloned pcGH gene was analysed. Then
recombinant pcGH was produced both in intracellular and periplasmic manner. And
the factors effecting the expression level of the Escherichia coli were discussed.
The newly cloned pcGH showed high homogeneity to pcGH and Sciaenops
ocellata GH from GenBank in different regions respectively. Therefore this gene was
presumed to be the hybrid of several species of fish. This supposition will contribute
to find out the conserved regions of fGH. It'll also help to investigate the influence of
genetic mutation of GH on the growth of Pseudosciaena crocea.
The pcGH gene was cloned into pET-22b(+) for intracellular expression.
Recombinant plasmid (pG2) were transformed into E. coli BL21(DE3) and
Rosetta(DE3). SDS-PAGE analysis showed that a specific protein band with the same
molecular weight as theoretical one of pcGH was induced in pG2/Rosetta(DE3); but
in the case of pG2/BL21(DE3), this band was difficult to be found. Therefore, it’s
concluded that Rosetta(DE3) can help to lighten the effect of rare codon on the
expression level in E. coli. The product of pG2/Rosetta(DE3) was proved soluble,
then it was purified to be electrophoretically pure by ammonium sulfate precipitation
and cation exchange chromatography.
Another pair of primer was designed to insert the gene of pcGH into the
II
downstream of pel B signal peptide in pET-22b(+). The recombinant plasmid was
transformed into E. coli Rosetta(DE3). With IPTG induction, part of the recombinant
pGH was secreted into the periplasm of transformants.
Factors effecting the expression level of pcGH in E. coli, both on transcription
level and translation level, were discussed integrallty and systemically. Approches to
improve the expression level were put forward .
These works may have potential value in the production and application of
pcGH.
引文
[1] T Sakamoto, BS Shepherd, SS Madsen, et al. Osmoregulatory actions of growth
hormone and prolactin in an advanced teleost. Gen Comp Endocrinol, 1997, 106(1):
95-101.
[2] P Prunet, G Boeuf, and LM Houdebine. Plasma and pituitary prolactin levels in
rainbow trout during adaptation to different salinities. J Exp Zool, 1985,
235(2): 187-96.
[3] Brian S. Shepherd, Tatsuya Sakamoto, et al. Somatotropic actions of the
homologous growth hormone and prolactins in the euryhaline teleost, the tilapia,
Oreochromis mossambicus. Physiology, 1997,94： 2068-2072.
[4] RC Fargher and BA McKeown. The effect of prolactin on calcium homeostasis in
coho salmon (Oncorhynchus kisutch). Gen Comp Endocrinol, 1989， 73(3): 398-403.
[5] MJ Herrero-Turrion, RE Rodriguez, A Velasco, et al. Differential expression and
cellular localization of somatolactin-1 and -2 during early development in the
gilthead sea bream. Gen Comp Endocrinol, 2003; 132(1): 77-87.
[6] M Olivereau and J Olivereau. Calcium-sensitive cells of the pars intermedia and
osmotic balance in the eel. I. Responses to changes in the environmental calcium
and magnesium. Cell Tissue Res, 1982, 222(2): 231-41.
[7] GJ Van Eys and SE Wendelaar Bonga. Responses of the PAS-positive pars intermedia
cells in the cichlid fish Sarotherodon mossambicus to ambient calcium and
background adaptation. Cell Tissue Res, 1984， 236(1): 181-7.
[8] CA Berghs and EW Roubos. Background adaptation and synapse plasticity in the
pars intermedia of Xenopus laevis. Neuroscience, 1996, 70(3): 833-41.
[9] GJ van Eys. Structural changes in the pars intermedia of the cichlid teleost
Sarotherodon mossambicus as a result of background adaptation and illumination.
II. The PAS-positive cells. Cell Tissue Res, 1980, 210(2): 171-9
[10] M Rand-Weaver, P Swanson, H Kawauchi, et al. Somatolactin, a novel pituitary
protein: purification and plasma levels during reproductive maturation of coho
salmon. J Endocrinol, 1992,133(3): 393-403.
[11] M Ankersen, TK Hansen, I Ahnfelt-Ronne I, et al. Growth hormone
secretagogues: recent advances and applications. Drug Discov Today, 1999; 4(11):
497-506.
[12] TA Marchant, JG Dulka, and RE Peter. Relationship between serum growth
hormone levels and the brain and pituitary content of immunoreactive
somatostatin in the goldfish, Carassius auratus L. Gen Comp Endocrinol, 1989;
73(3): 458-68.
49
福州大学硕士学位论文
[13] M Montero, L Yon, S Kikuyama, et al. Molecular evolution of the growth
hormone-releasing hormone/pituitary adenylate cyclase-activating polypeptide
gene family. Functional implication in the regulation of growth hormone
secretion. J Mol Endocrinol, 2000; 25(2): 157-68.
[14] JP Chang, H Cook, GL Freedman, et al. Use of a pituitary cell dispersion
method and primary culture system for the studies of gonadotropin-releasing
hormone action in the goldfish, Carassius auratus. I. Initial morphological,
static, and cell column perifusion studies. Gen Comp Endocrinol, 1990; 77(2):
256-73.
[15] P Melamed, N Eliahu, B Levavi-Sivan, et al. Hypothalamic and thyroidal
regulation of growth hormone in tilapia. Gen Comp Endocrinol, 1995; 97(1):
13-30.
[16] Blaise O, Le Bail P Y, Weil C. Lack of gonadotropin releasing hormone action
on in vivo and in vitro growth hormone release, in rainbow trout (Oncorhynchus
mykiss). Comp. Biochem. Physiol., 1995,110C:133-141.
[17] Bosma P T, Van Dijk W, Van Haren S F, et al,. GnRH receptors are restricted
to gonadotropes in male African catfish[A]. In: Goetz F W, Thomas P(eds.),
Reproductive physiology of fish[M]. Austin, U. S. A. : Fish Symposium,
1995,95:55-57.
[18] 林浩然. 神经内分泌因子调控鱼类生殖和生长的相互作用. 动物学研究. 2000,
21(1):12-16.
[19] V L Trudeau, G M Somoza, C S Nahorniak, et al. Interactions of estradiol
with gonadotropin-releasing hormone and thyrotropin-releasing hormone in the
control of growth hormone secretion in the goldfish. Neuroendocrinology, 1992,
56(4): 483-90.
[20] Farmer S.W., H.Papoff, T.Hayashida,et al. Isolation and properties of
teleostean growth hormone. Gen. Comp. Endocrinol.1976,30:91～100.
[21] 徐斌，张培军，李德尚。鱼类生长激素的分离、鉴定及其生理功能研究的进展。海
洋与湖泊。1997（2）：209～214.
[22] H Kawauchi, S Moriyama, A Yasuda, et al. Isolation and characterization of
chum salmon growth hormone. Arch Biochem Biophys, 1986, 244(2): 542-52.
[23] V Skibeli, O Andersen, and KM Gautvik. Purification and characterization
of Atlantic salmon growth hormone and evidence for charge heterogeneity. Gen
Comp Endocrinol, 1990, 80(3): 333-44.
[24] T Noso, A Yasuda, I Kawazoe, et al. Isolation and characterization of growth
hormone from a marine fish, bonito (Katsuwonus pelamis). Int J Pept Protein Res,
1988, 32(6): 579-89.
[25] K Watanabe, A Igarashi, T Noso, et al. Chemical identification of catfish
growth hormone and prolactin. Mol Mar Biol Biotechnol, 1992, 1(3): 239-49.
[26] LB Agellon and TT Chen. Rainbow trout growth hormone: molecular cloning of
cDNA and expression in Escherichia coli. DNA, 1986, 5(6): 463-71.
50
大黄鱼生长激素的胞内表达和周质表达
[27] B Johansen, OC Johnsen, and S Valla. The complete nucleotide sequence of
the growth-hormone gene from Atlantic salmon (Salmo salar). Gene, 1989, 77(2):
317-24.
[28] CS Chiou, HT Chen, and WC Chang. The complete nucleotide sequence of the
growth-hormone gene from the common carp (Cyprinus carpio). Biochim Biophys Acta,
1990, 1087(1): 91-4.
[29] Z Zhu, L He, and TT Chen. Primary-structural and evolutionary analyses of
the growth-hormone gene from grass carp (Ctenopharyngodon idellus). Eur. J.
Biochem., 1992, 207: 643 - 648.
[30] Y Koren, S Sarid, R Ber, et al. Carp growth hormone: molecular cloning and
sequencing of cDNA. Gene, 1989, 77(2): 309-15.
[31] V Skibeli, O Andersen, and KM Gautvik. Purification and characterization
of Atlantic salmon growth hormone and evidence for charge heterogeneity. Gen
Comp Endocrinol, 1990, 80(3): 333-44.
[32] SS Abdel-Meguid, HS Shieh, WW Smith, et al. Three-dimensional structure of
a genetically engineered variant of porcine growth hormone. 1987,84(18):
6434-7.
[33] AM de Vos, M Ultsch, and AA Kossiakoff. Human growth hormone and
extracellular domain of its receptor: crystal structure of the complex.
Science, 1992, 255(5042): 306-12.
[34] 孙逊，朱尚权. 生长激素的结构与功能. 国外医学生理、病理科学与临床分册，
1999,19(1)：6-9.
[35] P. W. Hochachka, T. P. Mommsen. 鱼类的生物化学与分子生物学.第二卷：分子
生物学前沿，（昌永华，余来宁译）.北京：中国农业出版社.2003.34-36.
[36] Jaw-Ching Liu, Kateryna D. Makova, Ronald M. Adkins, et al. Episodic
Evolution of Growth Hormone in Primates and Emergence of the Species Specificity
of Human Growth Hormone Receptor. Mol. Biol. Evol., 2001, 18: 945 - 953.
[37] 孙逊，朱尚权. 生长激素受体的结构、功能及其信号途径. 国外医学生理、病理科
学与临床分册 1999,19(1)：9－14.
[38] R Company, A Astola, C Pendon, et al. Somatotropic regulation of fish growth
and adiposity: growth hormone (GH) and somatolactin (SL) relationship. Comp
Biochem Physiol C Toxicol Pharmacol, 2001, 130(4): 435-45.
[39] HJ Tsai, KL Lin, JC Kuo, et al. Highly efficient expression of fish growth
hormone by Escherichia coli cells. Appl. Envir. Microbiol., 1995, 61: 4116 -
4119.
[40] 白俊杰,简清,马进，等。鲑鱼生长激素 cDNA 在大肠杆菌中的表达及表达产物对罗
非鱼促生长作用。农业生物技术学报 1998,6(4)：343-346.
[41] 马进，白俊杰，李新辉. 虹鳟生长激素 cDNA 在酵母中的表达。生物工程学报，1999，
15（4）：434～438.
51
福州大学硕士学位论文
[42] Jeffrey T. Silverstein, Wolters, William R., et al. Bovine growth hormone
treatment of channel catfish: strain and temperature effects on growth, plasma
IGF-I levels, feed intake and efficiency and body composition. Aquaculture
2000,190(1-2):77-88
[43] JR Markert, DA Higgs, HM Dye, et al. Influence of bovine growth hormone on
growth rate, appetite, and food conversion of yearling coho salmon (Oncorhynchus
kisutch) fed two diets of different composition. Can J Zool, 1977, 55(1): 74-83.
[44] Cook J T. Growth rate, body composition and feed digestibility/ convertion
of growth-enhanced transgenic Atlantic salmon (salmo salar). Aqua,
2000,188:15-32.
[45] 王伟，汪亚平，朱作言，等。酵母表达的重组草鱼生长激素对鲤鱼的促生长效应。
高技术通讯。2000（12）：5～8.
[46] Higgs D.A., E.M. Donaldson, H.M. Dye , et al. 1976. Influence of bovine
growth hormone and L-thyroxine on growth, muscle composition, and histological
structure of the gonads, thyroid, and pancreas and pituitary of coho salmon
(Oncorhynchus kisutch). J. Fish. Res. Board Can, 1976,33:1583-1603.
[47] JP Bolton, NL Collie, H Kawauchi, et al. Osmoregulatory actions of growth
hormone in rainbow trout (Salmo gairdneri). J Endocrinol, 1987, 112(1): 63-8.
[48] JM Shrimpton, RH Devlin, E McLean, et al. Increases in gill cytosolic
corticosteroid receptor abundance and saltwater tolerance in juvenile coho
salmon (Oncorhynchus kisutch) treated with growth hormone and placental
lactogen. Gen Comp Endocrinol. 1995, 98(1): 1-15.
[49] G. Flik, W. Atsma,, J. C. Fenwick, et al. Homologous recombinant growth
hormone and calcium metabolism in the tilapia, oreochromis mossambicus, adapted
to fresh water. Journal of Experimental Biology, 1993,185(1): 107-119
[50] G Van der Kraak, PM Rosenblum, and RE Peter. Growth hormone-dependent
potentiation of gonadotropin-stimulated steroid production by ovarian
follicles of the goldfish. Gen Comp Endocrinol, 1990, 79(2): 233-9.
[51] JA Calduch-Giner, A Sitjà-Bobadilla, P Alvarez-Pellitero, et al. Growth
hormone as an in vitro phagocyte-activating factor in the gilthead sea bream
(Sparus aurata). Cell Tissue Res, 1997, 287(3): 535-40.
[52] J Perez Sanchez, J Smal, and PY Le Bail. Location and characterization of
growth hormone binding sites in the central nervous system of a teleost fish
(Oncorhynchus mykiss). Growth Regul, 1991, 1(4): 145-52.
[53] E Jonsson, JI Johnsson, and BT Bjornsson. Growth hormone increases
aggressive behavior in juvenile rainbow trout. Horm Behav, 1998, 33(1): 9-15.
[54] AR Foster, DF Houlihan, C Gray, et al. The effects of ovine growth hormone
on protein turnover in rainbow trout. Gen Comp Endocrinol, 1991, 82(1): 111-20.
[55] 李英华，鲤鱼生长激素 cDNA 在毕赤酵母中的胞内表达和分泌表达：[硕士学位论文].
广州：暨南大学，2001.
52
大黄鱼生长激素的胞内表达和周质表达
[56] M Fine, E Sakal, D Vashdi,et al. Recombinant carp (Cyprinus carpio) growth
hormone: expression, purification, and determination of biological activity in
vitro and in vivo. Gen Comp Endocrinol, 1993,89(1): 51-61.
[57] 白俊杰、罗建仁.基因重组鱼生长激素的表达及促生长作用研究进展(综述)。见：
2000 年中国水产学会学术年会论文集 迈向 21 世纪的渔业科技创新. 海洋出版社,
2000: 70-74.
[58] A Bartke. Can growth hormone (GH) accelerate aging? Evidence from
GH-transgenic mice. Neuroendocrinology, 2003; 78(4): 210-6.
[59] Macari M, Carneiro DJ, Larson ML, et al.Influence of dietary protein intake
and recombinant human somatotropin administration on growth and body
composition of juvenile tacambu (Praractus mesopotamicus x Colossoma
macroponius cross). Aquaculture， 1994,127:363-369.
[60] 崔宗斌，朱作言，崔奕波,等. 转人生长激素基因红鲤 F2 代阳性鱼的摄食及代谢研
究. 科学通报，1995，40(16)：1514-1517.
[61] Fu C., Cui Y., Hung S. S. O.,et al. Growth and feed utilization by F4 human
growth hormone transgenic carp fed diets with different protein levels. Journal
of Fish Biology. 1998,53(1):115-129.
[62] Leena S.，Shameena B.，Oommen O.V. In vivo and in vitro effects of prolactin
and growth hormone on lipid metabolism in a teleost, Anabas testudineus (Bloch).
Comparative Biochemistry and Physiology, 2001,128(4B): 761-766.
[63] Y Inui, S Miwa, and H Ishioka. Effect of mammalian growth hormone on amino
nitrogen mobilization in the eel. Gen Comp Endocrinol, 1985, 59(2): 287-94.
[64] TC Leung, TB Ng, and NY Woo. Metabolic effects of bovine growth hormone in
the tilapia Oreochromis mossambicus. Comp Biochem Physiol A, 1991,99(4): 633-6.
[65] Ahsan,et al. Role of growth hormone in regulation of blood glucose and
glycogen content of an Indian teleost, Clarias batrachus (Linn.).
1984(9):178-182.
[66] 傅萃长,朱作言.生长激素及其基因转移对鱼类生长和渗透压的调节作用. 动物学
报. 2000,46(4):407-415.
[67] J Calduch-Giner , H Duval, F Chesnel, et al. Fish growth hormone receptor:
molecular characterization of two membrane-anchored forms. Endocrinology,
2001,142(7): 3269-73.
[68] L. S. Argetsinger and C. Carter-Su. Mechanism of signaling by growth hormone
receptor. Physiol Rev, 1996,76: 1089 - 1107.
[69] AM de Vos, M Ultsch, and AA Kossiakoff. Human growth hormone and
extracellular domain of its receptor: crystal structure of the complex. Science,
1992, 255(5042): 306-12.
[70] S Takeda and K Inoue. Growth hormone-binding protein. Nippon Rinsho, 1995，
53(Su Pt 2): 346-50.
53
福州大学硕士学位论文
[71] B Funkenstein, A Silbergeld, B Cavari, et al. Growth hormone increases
plasma levels of insulin-like growth factor (IGF-I) in a teleost, the gilthead
seabream (sparus aurata).J Endocrinol, 1989, 120(2): 19-21.
[72] ES Gray and KM Kelley. Growth regulation in the gobiid teleost, Gillichthys
mirabilis: roles of growth hormone, hepatic growth hormone receptors and
insulin-like growth factor-I. J Endocrinol, 1991,131(1): 57-66.
[73] SD McCormick, T Sakamoto, S Hasegawa,et al. Osmoregulatory actions of
insulin-like growth factor-I in rainbow trout (Oncorhynchus mykiss). J
Endocrinol, 1991, 130(1): 87-92.
[74] QP Cao, SJ Duguay, E Plisetskaya, et al. Nucleotide sequence and growth
hormone-regulated expression of salmon insulin-like growth factor I mRNA. Mol.
Endocrinol., Dec 1989; 3: 2005 - 2010.
[75] CM Duan and Y Inui. Effects of recombinant eel growth hormone on the uptake
of [35S]sulfate by ceratobranchial cartilages of the Japanese eel, Anguilla
japonica. Gen Comp Endocrinol, 1990,79(2): 320-5.
[76] SD McCormick, PI Tsai, KM Kelley, et al. Hormonal control of sulfate uptake
by branchial cartilage of coho salmon: role of IGF-I. J Exp Zool, 1992,262(2):
166-71.
[77] Bern. Insulin-like growth factors under water: role in growth and function
of fish and other poikilothermic vertebrates. 1991: 150-154.
[78] Bern H. A., S. D. McCormick, K. M. Kelly. Insulin-like growth factors under
water：role in growth and function of fish and other poikilothermic vertebrates.
In: McCormick, K. M. ed. Morden Concepts of Insulin-like Growth Factors. New
York: Elesevier. 150-154.
[79] SD McCormick, KM Kelley, G Young, et al. Stimulation of coho salmon growth
by insulin-like growth factor I. Gen Comp Endocrinol, 1992, 86(3): 398-406.
[80] Li G. H., Z. B. Cui and Z. Y. Zhu. The growth enhancement of
pCAsIGF-transgenic deepbody crucian carp (Carassius cuvievi). Annual Report of
FEBL.1995:139-142.
[81] JJ Nagler, SJ Hwang, and DR Idler. Isolation of sockeye salmon growth hormone
utilizing serum triiodothyronine enhancement in rainbow trout to monitor
biological activity. Gen Comp Endocrinol, 1991，84(3): 374-80.
[82] 陈松林，邓文涛，贺路，等. 用酶联免疫吸附受体法检测鱼类生长激素的生物活性.
1995，19（3）：217-224.
[83] 张培军，徐永立,王艳秋.鲑鱼生长激素基因表达产物的促生长效应.海洋科学，1995
（6）：1-3.
[84] Zhu, Z.,G. Li, L. He,et al. Novel gene transfer into the fertilized eggs
of goldfish (Carassius auratus). Z. Angew . Ichthyol. 1985 (1): 1-34.
[85] SJ Du, ZY Gong, GL Fletcher, et al. Growth enhancement in transgenic Atlantic
salmon by the use of an "all fish" chimeric growth hormone gene construct.
Biotechnology (N Y), 1992, 10(2): 176-81.
54
大黄鱼生长激素的胞内表达和周质表达
[86] 崔宗斌，谢岳峰，许克圣，等. 人生长激素和转移的人生长激素基因对去垂体鱼的
生长代偿效应. 水生生物学报, 1993，17(2):166-173.
[87] Chatakondi N, Lovell R T, Duncan P L , et al. Body composition of transgenic
common carp, Cyprinus carpio, containing rainbow trout growth hormone gene.
Aquaculture , 1995, 138(1-4):99-109.
[88] 赵浩斌, 陈尚萍, 朱作言. 转基因鱼的研究进展. 农业生物技术学报. 1999，7
（3）:301-306.
[89] Sekine S. Cloning and expression of cDNA for salmon growth hormone in
Escherchia coli. Proc. Natl. Acad. SciU. S. A, 1985,82:4306-4310.
[90] 赵晓祥，张淑梅，杨学成，等.鲤鱼生长激素基因的改造及其在大肠杆菌中的表达.
生物技术，1995，5(5):11-15.
[91] 简清，白俊杰，马进，等。饲料中添加重组鱼生长激素对罗非鱼鱼种的促生长作用
研究。淡水鱼业，1999，29（3）：3-5.
[92] 白俊杰,马进,简清,等.鲤鱼(Cyprinuscarpio)生长激素基因克隆及原核表达.中国
生物化学与分子生物学报，1999，15(3):409-412.
[93] 张同海，宋诗铎，赵为诚，等.鲑鱼生长激素基因分泌型表达质粒的构建.生物化学
与生物物理进展，1994，21(4):338-379.
[94] 何为,陈慧.大黄鱼增养殖的现状、存在的问题及发展对策. 2000 年中国水产学术
年会论文集，2000：345～349.
[95] 江树勋,马鸿媚，邓文汉，等. 大黄鱼生长激素基因的分离及序列测定. 生物工程
进展，2002, 22(2)：88-90.
[96] KJ Jeong and SY Lee. Secretory production of human granulocyte
colony-stimulating factor in Escherichia coli. Protein Expr Purif, 2001; 23(2):
311-8.
[97] Belin and PL Boquet. The Escherichia coli dsbA gene is partly transcribed
from the promoter of a weakly expressed upstream gene. Microbiology, 1994; 140:
3337 - 3348.
[98] CS Hoffman and A Wright. Fusions of secreted proteins to alkaline
phosphatase: an approach for studying protein secretion. PNAS, 1985; 82(15):
5107-11.
[99] S MacIntyre, R Freudl, M Degen, et al. The signal sequence of an Escherichia
coli outer membrane protein can mediate translocation of a not normally secreted
protein across the plasma membrane. J. Biol. Chem., 1987,262: 8416 - 8422.
[100] A. V. Zavialov, N. V. Batchikova, T. Korpela, et al. Secretion of Recombinant
Proteins via the Chaperone/Usher Pathway in Escherichia coli. Appl. Envir.
Microbiol., 2001,67(4): 1805 - 1814.
[101] Sato T., Matsui H., Shibahara S.,et al. New approaches for the high-level
expression of human interleukin-2 cDNA in Escherichia coli. J Biochem (Tokyo)，
1987，101: 525-534.
[102] Gold L.. Posttranscriptional regulatory mechanisms in Escherichia coli.
Annu. Rev. Biochem. 1988,57:199-233.
55
福州大学硕士学位论文
[103] 卢圣栋,等.现代分子生物学实验技术[Ｍ].北京:高等教育出版社,1993.
[104] 白俊杰,马进,简清,等。鱼生长激素基因在大肠杆菌中表达水平的研究。中国水产
科学。2000，7（1）：31-34.
[105] 吴加金,李伍举,王嘉玺.影响外源基因在原核中表达水平因素分析.生物工程学
报,1997,12(增刊):92-96.
[106] 李伍举，吴加金.RNA 二级结构的预测. 军事医学科学院院刊，1996，20(4):
298-301.
[107] 于长春，程焉平.蛋白质合成终止的研究进展.生命的化学，1996,16(4):27-29.
[108] H Grosjean and W Fiers. Preferential codon usage in prokaryotic genes: the
optimal codon-anticodon interaction energy and the selective codon usage in
efficiently expressed genes. Gene, June 1, 1982; 18(3): 199-209.
[109] R Schwartz and JF Curran. Analyses of frameshifting at UUU-pyrimidine sites.
Nucleic Acids Res., May 1997; 25: 2005 - 2011.
[110] T Wakagi, T Oshima, H Imamura, et al. Cloning of the gene for inorganic
pyrophosphatase from a thermoacidophilic archaeon, Sulfolobus sp. strain 7, and
overproduction of the enzyme by coexpression of tRNA for arginine rare codon.
Biosci Biotechnol Biochem, December 1, 1998; 62(12): 2408-14.
[111] BJ Del Tito, Jr, JM Ward, J Hodgson,et al. Effects of a minor isoleucyl tRNA
on heterologous protein translation in Escherichia coli. J. Bacteriol.,
1995,177(24):7086-7091.
[112] JF Kane. Effects of rare codon clusters on high-level expression of
heterologous proteins in Escherichia coli. Curr Opin Biotechnol, October 1, 1995;
6(5): 494-500.
[113] 杨喆，李太华。大肠杆菌中外源基因表达的研究进展。微生物学免疫学进展。2000，
28（2）：69-72.
[114] Miller K. W., Evans R. J., Eisenberg S. P., et al. Secretory leukocyte
protease inhibitor binding to mRNA and DNA as a possible cause of toxicity to
Escherichia coli. J. Bacteriol, 1989,171: 2166–2172.
hormone and prolactin in an advanced teleost. Gen Comp Endocrinol, 1997, 106(1):
95-101.
[2] P Prunet, G Boeuf, and LM Houdebine. Plasma and pituitary prolactin levels in
rainbow trout during adaptation to different salinities. J Exp Zool, 1985,
235(2): 187-96.
[3] Brian S. Shepherd, Tatsuya Sakamoto, et al. Somatotropic actions of the
homologous growth hormone and prolactins in the euryhaline teleost, the tilapia,
Oreochromis mossambicus. Physiology, 1997,94： 2068-2072.
[4] RC Fargher and BA McKeown. The effect of prolactin on calcium homeostasis in
coho salmon (Oncorhynchus kisutch). Gen Comp Endocrinol, 1989， 73(3): 398-403.
[5] MJ Herrero-Turrion, RE Rodriguez, A Velasco, et al. Differential expression and
cellular localization of somatolactin-1 and -2 during early development in the
gilthead sea bream. Gen Comp Endocrinol, 2003; 132(1): 77-87.
[6] M Olivereau and J Olivereau. Calcium-sensitive cells of the pars intermedia and
osmotic balance in the eel. I. Responses to changes in the environmental calcium
and magnesium. Cell Tissue Res, 1982, 222(2): 231-41.
[7] GJ Van Eys and SE Wendelaar Bonga. Responses of the PAS-positive pars intermedia
cells in the cichlid fish Sarotherodon mossambicus to ambient calcium and
background adaptation. Cell Tissue Res, 1984， 236(1): 181-7.
[8] CA Berghs and EW Roubos. Background adaptation and synapse plasticity in the
pars intermedia of Xenopus laevis. Neuroscience, 1996, 70(3): 833-41.
[9] GJ van Eys. Structural changes in the pars intermedia of the cichlid teleost
Sarotherodon mossambicus as a result of background adaptation and illumination.
II. The PAS-positive cells. Cell Tissue Res, 1980, 210(2): 171-9
[10] M Rand-Weaver, P Swanson, H Kawauchi, et al. Somatolactin, a novel pituitary
protein: purification and plasma levels during reproductive maturation of coho
salmon. J Endocrinol, 1992,133(3): 393-403.
[11] M Ankersen, TK Hansen, I Ahnfelt-Ronne I, et al. Growth hormone
secretagogues: recent advances and applications. Drug Discov Today, 1999; 4(11):
497-506.
[12] TA Marchant, JG Dulka, and RE Peter. Relationship between serum growth
hormone levels and the brain and pituitary content of immunoreactive
somatostatin in the goldfish, Carassius auratus L. Gen Comp Endocrinol, 1989;
73(3): 458-68.
49
福州大学硕士学位论文
[13] M Montero, L Yon, S Kikuyama, et al. Molecular evolution of the growth
hormone-releasing hormone/pituitary adenylate cyclase-activating polypeptide
gene family. Functional implication in the regulation of growth hormone
secretion. J Mol Endocrinol, 2000; 25(2): 157-68.
[14] JP Chang, H Cook, GL Freedman, et al. Use of a pituitary cell dispersion
method and primary culture system for the studies of gonadotropin-releasing
hormone action in the goldfish, Carassius auratus. I. Initial morphological,
static, and cell column perifusion studies. Gen Comp Endocrinol, 1990; 77(2):
256-73.
[15] P Melamed, N Eliahu, B Levavi-Sivan, et al. Hypothalamic and thyroidal
regulation of growth hormone in tilapia. Gen Comp Endocrinol, 1995; 97(1):
13-30.
[16] Blaise O, Le Bail P Y, Weil C. Lack of gonadotropin releasing hormone action
on in vivo and in vitro growth hormone release, in rainbow trout (Oncorhynchus
mykiss). Comp. Biochem. Physiol., 1995,110C:133-141.
[17] Bosma P T, Van Dijk W, Van Haren S F, et al,. GnRH receptors are restricted
to gonadotropes in male African catfish[A]. In: Goetz F W, Thomas P(eds.),
Reproductive physiology of fish[M]. Austin, U. S. A. : Fish Symposium,
1995,95:55-57.
[18] 林浩然. 神经内分泌因子调控鱼类生殖和生长的相互作用. 动物学研究. 2000,
21(1):12-16.
[19] V L Trudeau, G M Somoza, C S Nahorniak, et al. Interactions of estradiol
with gonadotropin-releasing hormone and thyrotropin-releasing hormone in the
control of growth hormone secretion in the goldfish. Neuroendocrinology, 1992,
56(4): 483-90.
[20] Farmer S.W., H.Papoff, T.Hayashida,et al. Isolation and properties of
teleostean growth hormone. Gen. Comp. Endocrinol.1976,30:91～100.
[21] 徐斌，张培军，李德尚。鱼类生长激素的分离、鉴定及其生理功能研究的进展。海
洋与湖泊。1997（2）：209～214.
[22] H Kawauchi, S Moriyama, A Yasuda, et al. Isolation and characterization of
chum salmon growth hormone. Arch Biochem Biophys, 1986, 244(2): 542-52.
[23] V Skibeli, O Andersen, and KM Gautvik. Purification and characterization
of Atlantic salmon growth hormone and evidence for charge heterogeneity. Gen
Comp Endocrinol, 1990, 80(3): 333-44.
[24] T Noso, A Yasuda, I Kawazoe, et al. Isolation and characterization of growth
hormone from a marine fish, bonito (Katsuwonus pelamis). Int J Pept Protein Res,
1988, 32(6): 579-89.
[25] K Watanabe, A Igarashi, T Noso, et al. Chemical identification of catfish
growth hormone and prolactin. Mol Mar Biol Biotechnol, 1992, 1(3): 239-49.
[26] LB Agellon and TT Chen. Rainbow trout growth hormone: molecular cloning of
cDNA and expression in Escherichia coli. DNA, 1986, 5(6): 463-71.
50
大黄鱼生长激素的胞内表达和周质表达
[27] B Johansen, OC Johnsen, and S Valla. The complete nucleotide sequence of
the growth-hormone gene from Atlantic salmon (Salmo salar). Gene, 1989, 77(2):
317-24.
[28] CS Chiou, HT Chen, and WC Chang. The complete nucleotide sequence of the
growth-hormone gene from the common carp (Cyprinus carpio). Biochim Biophys Acta,
1990, 1087(1): 91-4.
[29] Z Zhu, L He, and TT Chen. Primary-structural and evolutionary analyses of
the growth-hormone gene from grass carp (Ctenopharyngodon idellus). Eur. J.
Biochem., 1992, 207: 643 - 648.
[30] Y Koren, S Sarid, R Ber, et al. Carp growth hormone: molecular cloning and
sequencing of cDNA. Gene, 1989, 77(2): 309-15.
[31] V Skibeli, O Andersen, and KM Gautvik. Purification and characterization
of Atlantic salmon growth hormone and evidence for charge heterogeneity. Gen
Comp Endocrinol, 1990, 80(3): 333-44.
[32] SS Abdel-Meguid, HS Shieh, WW Smith, et al. Three-dimensional structure of
a genetically engineered variant of porcine growth hormone. 1987,84(18):
6434-7.
[33] AM de Vos, M Ultsch, and AA Kossiakoff. Human growth hormone and
extracellular domain of its receptor: crystal structure of the complex.
Science, 1992, 255(5042): 306-12.
[34] 孙逊，朱尚权. 生长激素的结构与功能. 国外医学生理、病理科学与临床分册，
1999,19(1)：6-9.
[35] P. W. Hochachka, T. P. Mommsen. 鱼类的生物化学与分子生物学.第二卷：分子
生物学前沿，（昌永华，余来宁译）.北京：中国农业出版社.2003.34-36.
[36] Jaw-Ching Liu, Kateryna D. Makova, Ronald M. Adkins, et al. Episodic
Evolution of Growth Hormone in Primates and Emergence of the Species Specificity
of Human Growth Hormone Receptor. Mol. Biol. Evol., 2001, 18: 945 - 953.
[37] 孙逊，朱尚权. 生长激素受体的结构、功能及其信号途径. 国外医学生理、病理科
学与临床分册 1999,19(1)：9－14.
[38] R Company, A Astola, C Pendon, et al. Somatotropic regulation of fish growth
and adiposity: growth hormone (GH) and somatolactin (SL) relationship. Comp
Biochem Physiol C Toxicol Pharmacol, 2001, 130(4): 435-45.
[39] HJ Tsai, KL Lin, JC Kuo, et al. Highly efficient expression of fish growth
hormone by Escherichia coli cells. Appl. Envir. Microbiol., 1995, 61: 4116 -
4119.
[40] 白俊杰,简清,马进，等。鲑鱼生长激素 cDNA 在大肠杆菌中的表达及表达产物对罗
非鱼促生长作用。农业生物技术学报 1998,6(4)：343-346.
[41] 马进，白俊杰，李新辉. 虹鳟生长激素 cDNA 在酵母中的表达。生物工程学报，1999，
15（4）：434～438.
51
福州大学硕士学位论文
[42] Jeffrey T. Silverstein, Wolters, William R., et al. Bovine growth hormone
treatment of channel catfish: strain and temperature effects on growth, plasma
IGF-I levels, feed intake and efficiency and body composition. Aquaculture
2000,190(1-2):77-88
[43] JR Markert, DA Higgs, HM Dye, et al. Influence of bovine growth hormone on
growth rate, appetite, and food conversion of yearling coho salmon (Oncorhynchus
kisutch) fed two diets of different composition. Can J Zool, 1977, 55(1): 74-83.
[44] Cook J T. Growth rate, body composition and feed digestibility/ convertion
of growth-enhanced transgenic Atlantic salmon (salmo salar). Aqua,
2000,188:15-32.
[45] 王伟，汪亚平，朱作言，等。酵母表达的重组草鱼生长激素对鲤鱼的促生长效应。
高技术通讯。2000（12）：5～8.
[46] Higgs D.A., E.M. Donaldson, H.M. Dye , et al. 1976. Influence of bovine
growth hormone and L-thyroxine on growth, muscle composition, and histological
structure of the gonads, thyroid, and pancreas and pituitary of coho salmon
(Oncorhynchus kisutch). J. Fish. Res. Board Can, 1976,33:1583-1603.
[47] JP Bolton, NL Collie, H Kawauchi, et al. Osmoregulatory actions of growth
hormone in rainbow trout (Salmo gairdneri). J Endocrinol, 1987, 112(1): 63-8.
[48] JM Shrimpton, RH Devlin, E McLean, et al. Increases in gill cytosolic
corticosteroid receptor abundance and saltwater tolerance in juvenile coho
salmon (Oncorhynchus kisutch) treated with growth hormone and placental
lactogen. Gen Comp Endocrinol. 1995, 98(1): 1-15.
[49] G. Flik, W. Atsma,, J. C. Fenwick, et al. Homologous recombinant growth
hormone and calcium metabolism in the tilapia, oreochromis mossambicus, adapted
to fresh water. Journal of Experimental Biology, 1993,185(1): 107-119
[50] G Van der Kraak, PM Rosenblum, and RE Peter. Growth hormone-dependent
potentiation of gonadotropin-stimulated steroid production by ovarian
follicles of the goldfish. Gen Comp Endocrinol, 1990, 79(2): 233-9.
[51] JA Calduch-Giner, A Sitjà-Bobadilla, P Alvarez-Pellitero, et al. Growth
hormone as an in vitro phagocyte-activating factor in the gilthead sea bream
(Sparus aurata). Cell Tissue Res, 1997, 287(3): 535-40.
[52] J Perez Sanchez, J Smal, and PY Le Bail. Location and characterization of
growth hormone binding sites in the central nervous system of a teleost fish
(Oncorhynchus mykiss). Growth Regul, 1991, 1(4): 145-52.
[53] E Jonsson, JI Johnsson, and BT Bjornsson. Growth hormone increases
aggressive behavior in juvenile rainbow trout. Horm Behav, 1998, 33(1): 9-15.
[54] AR Foster, DF Houlihan, C Gray, et al. The effects of ovine growth hormone
on protein turnover in rainbow trout. Gen Comp Endocrinol, 1991, 82(1): 111-20.
[55] 李英华，鲤鱼生长激素 cDNA 在毕赤酵母中的胞内表达和分泌表达：[硕士学位论文].
广州：暨南大学，2001.
52
大黄鱼生长激素的胞内表达和周质表达
[56] M Fine, E Sakal, D Vashdi,et al. Recombinant carp (Cyprinus carpio) growth
hormone: expression, purification, and determination of biological activity in
vitro and in vivo. Gen Comp Endocrinol, 1993,89(1): 51-61.
[57] 白俊杰、罗建仁.基因重组鱼生长激素的表达及促生长作用研究进展(综述)。见：
2000 年中国水产学会学术年会论文集 迈向 21 世纪的渔业科技创新. 海洋出版社,
2000: 70-74.
[58] A Bartke. Can growth hormone (GH) accelerate aging? Evidence from
GH-transgenic mice. Neuroendocrinology, 2003; 78(4): 210-6.
[59] Macari M, Carneiro DJ, Larson ML, et al.Influence of dietary protein intake
and recombinant human somatotropin administration on growth and body
composition of juvenile tacambu (Praractus mesopotamicus x Colossoma
macroponius cross). Aquaculture， 1994,127:363-369.
[60] 崔宗斌，朱作言，崔奕波,等. 转人生长激素基因红鲤 F2 代阳性鱼的摄食及代谢研
究. 科学通报，1995，40(16)：1514-1517.
[61] Fu C., Cui Y., Hung S. S. O.,et al. Growth and feed utilization by F4 human
growth hormone transgenic carp fed diets with different protein levels. Journal
of Fish Biology. 1998,53(1):115-129.
[62] Leena S.，Shameena B.，Oommen O.V. In vivo and in vitro effects of prolactin
and growth hormone on lipid metabolism in a teleost, Anabas testudineus (Bloch).
Comparative Biochemistry and Physiology, 2001,128(4B): 761-766.
[63] Y Inui, S Miwa, and H Ishioka. Effect of mammalian growth hormone on amino
nitrogen mobilization in the eel. Gen Comp Endocrinol, 1985, 59(2): 287-94.
[64] TC Leung, TB Ng, and NY Woo. Metabolic effects of bovine growth hormone in
the tilapia Oreochromis mossambicus. Comp Biochem Physiol A, 1991,99(4): 633-6.
[65] Ahsan,et al. Role of growth hormone in regulation of blood glucose and
glycogen content of an Indian teleost, Clarias batrachus (Linn.).
1984(9):178-182.
[66] 傅萃长,朱作言.生长激素及其基因转移对鱼类生长和渗透压的调节作用. 动物学
报. 2000,46(4):407-415.
[67] J Calduch-Giner , H Duval, F Chesnel, et al. Fish growth hormone receptor:
molecular characterization of two membrane-anchored forms. Endocrinology,
2001,142(7): 3269-73.
[68] L. S. Argetsinger and C. Carter-Su. Mechanism of signaling by growth hormone
receptor. Physiol Rev, 1996,76: 1089 - 1107.
[69] AM de Vos, M Ultsch, and AA Kossiakoff. Human growth hormone and
extracellular domain of its receptor: crystal structure of the complex. Science,
1992, 255(5042): 306-12.
[70] S Takeda and K Inoue. Growth hormone-binding protein. Nippon Rinsho, 1995，
53(Su Pt 2): 346-50.
53
福州大学硕士学位论文
[71] B Funkenstein, A Silbergeld, B Cavari, et al. Growth hormone increases
plasma levels of insulin-like growth factor (IGF-I) in a teleost, the gilthead
seabream (sparus aurata).J Endocrinol, 1989, 120(2): 19-21.
[72] ES Gray and KM Kelley. Growth regulation in the gobiid teleost, Gillichthys
mirabilis: roles of growth hormone, hepatic growth hormone receptors and
insulin-like growth factor-I. J Endocrinol, 1991,131(1): 57-66.
[73] SD McCormick, T Sakamoto, S Hasegawa,et al. Osmoregulatory actions of
insulin-like growth factor-I in rainbow trout (Oncorhynchus mykiss). J
Endocrinol, 1991, 130(1): 87-92.
[74] QP Cao, SJ Duguay, E Plisetskaya, et al. Nucleotide sequence and growth
hormone-regulated expression of salmon insulin-like growth factor I mRNA. Mol.
Endocrinol., Dec 1989; 3: 2005 - 2010.
[75] CM Duan and Y Inui. Effects of recombinant eel growth hormone on the uptake
of [35S]sulfate by ceratobranchial cartilages of the Japanese eel, Anguilla
japonica. Gen Comp Endocrinol, 1990,79(2): 320-5.
[76] SD McCormick, PI Tsai, KM Kelley, et al. Hormonal control of sulfate uptake
by branchial cartilage of coho salmon: role of IGF-I. J Exp Zool, 1992,262(2):
166-71.
[77] Bern. Insulin-like growth factors under water: role in growth and function
of fish and other poikilothermic vertebrates. 1991: 150-154.
[78] Bern H. A., S. D. McCormick, K. M. Kelly. Insulin-like growth factors under
water：role in growth and function of fish and other poikilothermic vertebrates.
In: McCormick, K. M. ed. Morden Concepts of Insulin-like Growth Factors. New
York: Elesevier. 150-154.
[79] SD McCormick, KM Kelley, G Young, et al. Stimulation of coho salmon growth
by insulin-like growth factor I. Gen Comp Endocrinol, 1992, 86(3): 398-406.
[80] Li G. H., Z. B. Cui and Z. Y. Zhu. The growth enhancement of
pCAsIGF-transgenic deepbody crucian carp (Carassius cuvievi). Annual Report of
FEBL.1995:139-142.
[81] JJ Nagler, SJ Hwang, and DR Idler. Isolation of sockeye salmon growth hormone
utilizing serum triiodothyronine enhancement in rainbow trout to monitor
biological activity. Gen Comp Endocrinol, 1991，84(3): 374-80.
[82] 陈松林，邓文涛，贺路，等. 用酶联免疫吸附受体法检测鱼类生长激素的生物活性.
1995，19（3）：217-224.
[83] 张培军，徐永立,王艳秋.鲑鱼生长激素基因表达产物的促生长效应.海洋科学，1995
（6）：1-3.
[84] Zhu, Z.,G. Li, L. He,et al. Novel gene transfer into the fertilized eggs
of goldfish (Carassius auratus). Z. Angew . Ichthyol. 1985 (1): 1-34.
[85] SJ Du, ZY Gong, GL Fletcher, et al. Growth enhancement in transgenic Atlantic
salmon by the use of an "all fish" chimeric growth hormone gene construct.
Biotechnology (N Y), 1992, 10(2): 176-81.
54
大黄鱼生长激素的胞内表达和周质表达
[86] 崔宗斌，谢岳峰，许克圣，等. 人生长激素和转移的人生长激素基因对去垂体鱼的
生长代偿效应. 水生生物学报, 1993，17(2):166-173.
[87] Chatakondi N, Lovell R T, Duncan P L , et al. Body composition of transgenic
common carp, Cyprinus carpio, containing rainbow trout growth hormone gene.
Aquaculture , 1995, 138(1-4):99-109.
[88] 赵浩斌, 陈尚萍, 朱作言. 转基因鱼的研究进展. 农业生物技术学报. 1999，7
（3）:301-306.
[89] Sekine S. Cloning and expression of cDNA for salmon growth hormone in
Escherchia coli. Proc. Natl. Acad. SciU. S. A, 1985,82:4306-4310.
[90] 赵晓祥，张淑梅，杨学成，等.鲤鱼生长激素基因的改造及其在大肠杆菌中的表达.
生物技术，1995，5(5):11-15.
[91] 简清，白俊杰，马进，等。饲料中添加重组鱼生长激素对罗非鱼鱼种的促生长作用
研究。淡水鱼业，1999，29（3）：3-5.
[92] 白俊杰,马进,简清,等.鲤鱼(Cyprinuscarpio)生长激素基因克隆及原核表达.中国
生物化学与分子生物学报，1999，15(3):409-412.
[93] 张同海，宋诗铎，赵为诚，等.鲑鱼生长激素基因分泌型表达质粒的构建.生物化学
与生物物理进展，1994，21(4):338-379.
[94] 何为,陈慧.大黄鱼增养殖的现状、存在的问题及发展对策. 2000 年中国水产学术
年会论文集，2000：345～349.
[95] 江树勋,马鸿媚，邓文汉，等. 大黄鱼生长激素基因的分离及序列测定. 生物工程
进展，2002, 22(2)：88-90.
[96] KJ Jeong and SY Lee. Secretory production of human granulocyte
colony-stimulating factor in Escherichia coli. Protein Expr Purif, 2001; 23(2):
311-8.
[97] Belin and PL Boquet. The Escherichia coli dsbA gene is partly transcribed
from the promoter of a weakly expressed upstream gene. Microbiology, 1994; 140:
3337 - 3348.
[98] CS Hoffman and A Wright. Fusions of secreted proteins to alkaline
phosphatase: an approach for studying protein secretion. PNAS, 1985; 82(15):
5107-11.
[99] S MacIntyre, R Freudl, M Degen, et al. The signal sequence of an Escherichia
coli outer membrane protein can mediate translocation of a not normally secreted
protein across the plasma membrane. J. Biol. Chem., 1987,262: 8416 - 8422.
[100] A. V. Zavialov, N. V. Batchikova, T. Korpela, et al. Secretion of Recombinant
Proteins via the Chaperone/Usher Pathway in Escherichia coli. Appl. Envir.
Microbiol., 2001,67(4): 1805 - 1814.
[101] Sato T., Matsui H., Shibahara S.,et al. New approaches for the high-level
expression of human interleukin-2 cDNA in Escherichia coli. J Biochem (Tokyo)，
1987，101: 525-534.
[102] Gold L.. Posttranscriptional regulatory mechanisms in Escherichia coli.
Annu. Rev. Biochem. 1988,57:199-233.
55
福州大学硕士学位论文
[103] 卢圣栋,等.现代分子生物学实验技术[Ｍ].北京:高等教育出版社,1993.
[104] 白俊杰,马进,简清,等。鱼生长激素基因在大肠杆菌中表达水平的研究。中国水产
科学。2000，7（1）：31-34.
[105] 吴加金,李伍举,王嘉玺.影响外源基因在原核中表达水平因素分析.生物工程学
报,1997,12(增刊):92-96.
[106] 李伍举，吴加金.RNA 二级结构的预测. 军事医学科学院院刊，1996，20(4):
298-301.
[107] 于长春，程焉平.蛋白质合成终止的研究进展.生命的化学，1996,16(4):27-29.
[108] H Grosjean and W Fiers. Preferential codon usage in prokaryotic genes: the
optimal codon-anticodon interaction energy and the selective codon usage in
efficiently expressed genes. Gene, June 1, 1982; 18(3): 199-209.
[109] R Schwartz and JF Curran. Analyses of frameshifting at UUU-pyrimidine sites.
Nucleic Acids Res., May 1997; 25: 2005 - 2011.
[110] T Wakagi, T Oshima, H Imamura, et al. Cloning of the gene for inorganic
pyrophosphatase from a thermoacidophilic archaeon, Sulfolobus sp. strain 7, and
overproduction of the enzyme by coexpression of tRNA for arginine rare codon.
Biosci Biotechnol Biochem, December 1, 1998; 62(12): 2408-14.
[111] BJ Del Tito, Jr, JM Ward, J Hodgson,et al. Effects of a minor isoleucyl tRNA
on heterologous protein translation in Escherichia coli. J. Bacteriol.,
1995,177(24):7086-7091.
[112] JF Kane. Effects of rare codon clusters on high-level expression of
heterologous proteins in Escherichia coli. Curr Opin Biotechnol, October 1, 1995;
6(5): 494-500.
[113] 杨喆，李太华。大肠杆菌中外源基因表达的研究进展。微生物学免疫学进展。2000，
28（2）：69-72.
[114] Miller K. W., Evans R. J., Eisenberg S. P., et al. Secretory leukocyte
protease inhibitor binding to mRNA and DNA as a possible cause of toxicity to
Escherichia coli. J. Bacteriol, 1989,171: 2166–2172.