柞蚕核型多角体病毒lef-3基因、lef-1基因簇的研究及嗜酸性α-淀粉酶在家蚕中的表达
|
摘要
用随机克隆法,建立柞蚕核型多角体病毒(Antheraea pernyi nucleopolyhedrovirus,ApNPV)的质粒基因组文库:通过对插入片段进行克隆鉴定和序列分析,首次获得ApNPV的晚期表达因子3(later expression factor 3,lef-3)基因、蜕皮甾体尿苷二磷酸葡萄糖基转移酶基因(ecdysteroid UDP-glucosyltransferase,egt)、晚期表达因子1(lef-1)等基因,对其进行同源性比对分析。克隆得到了ApNPV PstI和XhoI两个片段,其中一个片断经序列分析表明:该片段含有完整的lef-3基因,与其它来源的lef-3基因具有很高的同源性。ApNPV lef-3基因的阅读框为1110个核苷酸,编码369个氨基酸,编码一种DNA单链结合蛋白(single-stranded DNA-binding protein,SSB)。在lef-3基因上游的互补序列有一编码127个氨基酸序列的不完全的阅读框。
第二个片段总长度为3388 bp,经序列分析表明:该片段包括5个完整读码框,与苜蓿银纹夜蛾多粒包埋核型多角体病毒(Autographa californica nucleopolyhedrovirus,AcNPV)ORF13/OpNPV ORF12同源性较高的基因、晚期表达因子1(later expression factor 1,lef-1)、缺失的Ecdysteroid UDP-Glucosyltransferase基因(egt)、da26、da16。其中egt基因编码蜕皮甾体尿苷二磷酸葡萄糖基转移酶;lef-1编码的是晚期表达因子1(LEF-1),分析得到LEF-1的氨基酸序列有3个高度保守同源区,具有引发酶的活性:da26基因编码结构蛋白BV/ODV-E26,这个蛋白在出芽型病毒(budded virus,BV)和包埋型病毒(occlusion-derived virus,ODV)中都存在。
本论文首次应用家蚕-杆状病毒表达系统表达嗜酸性α-淀粉酶,此酶基因长3906 bp,编码1301个氨基酸,蛋白质理论分子量140 KD。将克隆进大肠杆菌表达载体pET-22b(+)的amy基因克隆到转移载体pVL1393中,用常规方法获得病毒,感染家蚕后嗜酸性α-淀粉酶得到正确表达,检测表达产物具有淀粉酶活性。
The gene encoding single-stranded DNA-binding protein(SSB) late expression factor 3 was cloned from Pst I to Xho I fragment of Antheraea pemyi nucleopolyhedrovirus (ApNPV) genome, which is fisrt found in ApNPV. Nucleotide sequence analysis of fragment revealed that lef-3 is a whole open reading frame of 1110 bp codes for 369 amino acids. Compared to the LEF-3 from other baculovirus, it was shown that the homologous on amino acid level was high. LEF-3 has identified as being essential for baculovirus DNA replication. LEF-3 proteins contain a conserved pattern of amino acids similar to this SSB motif in their amino-terminal regions, indicating that this region may be important for LEF-3 function. Upstream the lef-3 there is a sequence encoded 127 amino acids, but it is a part open reading frame. According to analysis of homologous this nucleotide sequence is similar to open reading frame 73 (ORF) of Orgyia pseudotsugata nucleopolyhedrovirus (OpNPV). At 3'end there is no homologous sequence compared to OpNPV and Choristoneura fumiferana nucleopolyhedrovirus (CfNPV), which indicates particularity of ApNPV.We also fisrt found the gene encoding protein of similar to Autographa californica nucleopolyhedrovirus (AcNPV) ORF 13 ORF 12, LEF-1, truncated Ecdysteroid UDP-glucosyltransferase (EGT), BV/ODV-E26, DA 16, which were also cloned from another Pst I to Xho I fragment of ApNPV genome. Compared to the five genes from other baculovirus, they were shown that the homologous on amino acid level was high. Among the five genes: lef-1 is believed to encode LEF-1. LEF-1 has identified as being essential for baculovirus DNA replication and contains three conserved pattern of amino acids, indicating that these region may be important for LEF-1 function and LEF-1 also has the activity of primase. Downstream the lef-1 there is a sequence encoded 79 amino acids named egt, which encodes EGT. Compared to egt from other baculovirus, it is deficient. da26 encodes BV/ODV-E26, which is the structure protein of budded virus, (BV) and occlusion-derived virus(ODV).The α-amylase from the Gram-positive Alicylobacillus acidocaldarius was one kind of thermoacidophilic enzyme. It was expressed by Bombyx mori NPV expression vector fisrt. The nucleotide sequence of the gene amy was 3 906bp, comprising one open reading frame encoding a polypeptide of 1 301 amino acids, the calculated molecular weight of the a-amylase AMY was about 140KD. AMY was expressed in silkworm larva by injection with the recombinant virus. The expressed products showed amylase activity.
第二个片段总长度为3388 bp,经序列分析表明:该片段包括5个完整读码框,与苜蓿银纹夜蛾多粒包埋核型多角体病毒(Autographa californica nucleopolyhedrovirus,AcNPV)ORF13/OpNPV ORF12同源性较高的基因、晚期表达因子1(later expression factor 1,lef-1)、缺失的Ecdysteroid UDP-Glucosyltransferase基因(egt)、da26、da16。其中egt基因编码蜕皮甾体尿苷二磷酸葡萄糖基转移酶;lef-1编码的是晚期表达因子1(LEF-1),分析得到LEF-1的氨基酸序列有3个高度保守同源区,具有引发酶的活性:da26基因编码结构蛋白BV/ODV-E26,这个蛋白在出芽型病毒(budded virus,BV)和包埋型病毒(occlusion-derived virus,ODV)中都存在。
本论文首次应用家蚕-杆状病毒表达系统表达嗜酸性α-淀粉酶,此酶基因长3906 bp,编码1301个氨基酸,蛋白质理论分子量140 KD。将克隆进大肠杆菌表达载体pET-22b(+)的amy基因克隆到转移载体pVL1393中,用常规方法获得病毒,感染家蚕后嗜酸性α-淀粉酶得到正确表达,检测表达产物具有淀粉酶活性。
The gene encoding single-stranded DNA-binding protein(SSB) late expression factor 3 was cloned from Pst I to Xho I fragment of Antheraea pemyi nucleopolyhedrovirus (ApNPV) genome, which is fisrt found in ApNPV. Nucleotide sequence analysis of fragment revealed that lef-3 is a whole open reading frame of 1110 bp codes for 369 amino acids. Compared to the LEF-3 from other baculovirus, it was shown that the homologous on amino acid level was high. LEF-3 has identified as being essential for baculovirus DNA replication. LEF-3 proteins contain a conserved pattern of amino acids similar to this SSB motif in their amino-terminal regions, indicating that this region may be important for LEF-3 function. Upstream the lef-3 there is a sequence encoded 127 amino acids, but it is a part open reading frame. According to analysis of homologous this nucleotide sequence is similar to open reading frame 73 (ORF) of Orgyia pseudotsugata nucleopolyhedrovirus (OpNPV). At 3'end there is no homologous sequence compared to OpNPV and Choristoneura fumiferana nucleopolyhedrovirus (CfNPV), which indicates particularity of ApNPV.We also fisrt found the gene encoding protein of similar to Autographa californica nucleopolyhedrovirus (AcNPV) ORF 13 ORF 12, LEF-1, truncated Ecdysteroid UDP-glucosyltransferase (EGT), BV/ODV-E26, DA 16, which were also cloned from another Pst I to Xho I fragment of ApNPV genome. Compared to the five genes from other baculovirus, they were shown that the homologous on amino acid level was high. Among the five genes: lef-1 is believed to encode LEF-1. LEF-1 has identified as being essential for baculovirus DNA replication and contains three conserved pattern of amino acids, indicating that these region may be important for LEF-1 function and LEF-1 also has the activity of primase. Downstream the lef-1 there is a sequence encoded 79 amino acids named egt, which encodes EGT. Compared to egt from other baculovirus, it is deficient. da26 encodes BV/ODV-E26, which is the structure protein of budded virus, (BV) and occlusion-derived virus(ODV).The α-amylase from the Gram-positive Alicylobacillus acidocaldarius was one kind of thermoacidophilic enzyme. It was expressed by Bombyx mori NPV expression vector fisrt. The nucleotide sequence of the gene amy was 3 906bp, comprising one open reading frame encoding a polypeptide of 1 301 amino acids, the calculated molecular weight of the a-amylase AMY was about 140KD. AMY was expressed in silkworm larva by injection with the recombinant virus. The expressed products showed amylase activity.
引文
1.吕鸿声 昆虫病毒分子生物学(M),北京:农业科技出版社,1998
2.储瑞银,必怡德,吕鸿声,李载平,吴祥甫,乙型肝炎病毒表面抗原基因(pres2-S)在家蚕培养细胞中的表达,生物化学和生物物理学报,1990,22:385-389
3.洪华珠,杨红,中国生物防治,1995,11(2):84-88
4.李镇,龙綮新,袁美妗,贺雄雷,庞义,王珣章,斜纹夜蛾核多角体病毒(SpltNPV)基因组EcoR I-G片段的序列分析,中国病毒学,2002,17(2):119-124
5.林建城,酶在食品工业、轻工业和环境保护上的应用分析,莆田学院学报,2005,12(2):17-20
6.沈兴家,BmNPV和AcNPV egt基因启动子特性及家蚕海藻糖基因启动子活性的滞育激素调节研究[博士学位论文],北京:中国农业科学院研究生院,2004
7.石生林,柞蚕核型多角体病毒B-ORF6L、ptp、lef-12基因克隆及表达[博士学位论文],北京:中国科学院,2005
8.王海燕,秦浚川,王敖全,唐国敏,黑曲霉酸性α-淀粉酶基因和糖化酶基因对工业酒精酵母的整合及其共表达,微生物学报,2004,44(4):483-486
9.王文兵,张志芳,何家禄,吕鸿声,杆状病毒基冈组DNA复制基因的研究进展,生物化学与生物物理进展,2000,27(3):257-261
10.袁铁铮,姚斌,罗会颖,王亚茹,伍宁丰,范云六,来源于嗜热脂环酸杆菌的嗜酸性α-淀粉酶的研究,生物工程学报,2005,21(1):78-83
11.张春发,刘淑珊,范琦,李广泽,柞蚕核型多角体病毒载体表达系统基因工程研究进展,华南农业大学学报,1995,16(1):133-137
12.张纪忠,微生物分类学,上海:复旦大学出版社,1990,377-412
13.张丽苹,徐岩,金建中,酸性α-淀粉酶的研究与应用,酿酒,2002,29(3):19—21
14.张小霞,梁振普,牛国栋,等,棉铃虫核多角体病毒lef-3基因的序列分析,中国病毒学,2002,17(3):234—238
15.张志芳,何家禄,吴祥甫,家蚕核多角体病毒DNA聚合酶基因的克隆,蚕业科学,1995,21(2):86-90
16.张志芳,张颖,吕鸿声,等,家蚕核多角体病毒解链酶基因的克隆及部分序列分析,病毒学报,1994,10(4):381-383
17.张志芳,张颖,吕鸿声等,家蚕核多角体病毒DNA复制起始点hr3的结构功,中国科学B辑,1995,25(9):949-955
18. Ahrens CH, Rohrmann GF, Replication of Orgyia pseudotsugata baculovirus DNA: lef-2 and ie-1 are essential and ie-2, p34, and Op-iap are stimulatory genes, Virology, 1995, 212(2): 650-662
19. Ahrens CH, Russell R LQ, Funk CJ, The Sequence of the Orgyia pseudotsugata Multinucleocapsid Nuclear Polyhedrosis Virus Genome, Virology, 1997, 229:381-399
20. Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ, Basic local alignment search tool, J Mol Biol, 1990, 215:403-410
21. Antje C, Springer Handbook of Enzymes, 2nded, New York: Springer-Verlag Berlin Heidelberg, 2003, 1-39
22. Augustin M, R Huber, J Kaiser, Crystal structure of a DNA dependent RNA polymerase(DNA primase), Nat Struct Biol, 2001, 8:57-61
23. Barrett JW, Krell PJ, Arif BM, Characterization, sequencing and phylogeny of the ecdysteroid UDP-glucosyltransferase gene from two distinct nuclear polyhedrosis viruses isolated from Choristoneurafumiferana, J Gen Virol, 1995, 76:2447-2456
24. Blissard GW, Quant-Russell RL, Rohrmann GF, Beaudreau GS, Nucleotide Sequence, Transcriptional Mapping, and temporal expression of the gene encoding p39, a major structural protein of the muticapisid Nuclear Polyhedrosis Virus of Orgyia pseudotsugata, Virology, 1989, 168:354-362
25. Boyce FM, Bucher NL, Baculovirus-mediated gene transfer into mammalian cells, Proc Natl Acad Sci U S A, 1996, 93:2348-2352
26. Buonocore V, Caporale C, De Rosa M, Gambacorta A, Stable, inducible thermoacidophilic alpha-amylase from Bacillus acidocaldarius, J Bacteriol, 1976, 128(2):515-21
27. Carstens EB, Lu AL, Chan HB, Sequence, transcriptional mapping, and over expression of p47, a baculovirus gene regulating late gene expression, J Viro, 1993, 67:2513-2520
28. Chert XW, IJkel WF, Tarchini R, XL Sun, Sandbrink H, H Wang, Peters S, Zuidema D, Lankhorst RK, Vlak JM and ZH H, The sequence of the Helicoverpa armigera single nucleocapsid nucleopolyhedrovirus genome, J Gen Virol, 2001, 82:241-257
29. Christian HA, Cheryl Carlson, George FR, Identification, Sequence, and Transcriptional Analysis of lef-3, a Gene Essential for Orgyia pseudotsugata Baculovirus DNA Replication(J). Virology, 1995,210:372-382
30. Clarke EE, Tristem M, Cory JS, O'Reilly DR, Characterization of the ecdysteroid UDP-glucosyltransferase gene from Mamestra brassicae nucleopolyhedrovirus, J Gen Virol, 1996, 77:2865-2871
31. Clem RJ, Fechheimer M, Miller LK, Prevention of apoptosis by a baculovirus gene during infection of insect cells, Science, 1991,254:1388-1390
32. Copeland W, X Tan, Active site mapping of the catalytic mouse primase subunit by alanine scanning mutagenesis, J Biol Chem, 1995,270:3905-3913
33. Christian H A,Cheryl Carlson, George F R, Identification, Sequence, and Trans- criptional Analysis of lef-3, a Gene Essential for Orgyia pseudotsugata Baculovirus DNA Replication, Virology, 1995, 210:372-382
34. Copeland W, X Tan, Active site mapping of the catalytic mouse primase subunit by alanine scanning mutagenesis, J Biol Chem, 1995, 270:3905-3913
35. Dingwall C, Laskey RA, Nuclear targeting sequences-A consensus? TIBS 1991, 16:478-481
36. Evans JT, Leisy D J, Rohrmann GF, Characterization of the interaction between the baculovirus replication factors LEF-1 and LEF-2, J Virol, 1997, 71(4): 3114-3119
37. Flipsen JTM, Deletion of the baculovirus ecdysteroid UDP-glucosyltransferase gene induces early degeneration of Malpighian tubules on infected insects, J Virol, 1995, 69:4529-4532
38. Friesen PD, Miller LK, The regulation of baculovirus gene expression, Curr Top Microbiol Immunol, 1986, 131:31-49
39. Fukusumi, S, A Kamizono, S Horinouchi, T Beppu, Cloning and nucleotide sequence of a heat-stable amylase gene from an anaerobic thermophile, Dictyoglomus thermophilum, Eur J Biochem, 1988, 174:15-21
40. Gomi S, Majima K, Maeda S, Sequence analysis of the genome of Bombyx mori nucleopolyhedrovirus, J Gen Virol, 1999, 80(5): 1323-1337
41. Guoqlang Dong, Clatre Vieille, Alexei Savchenk, J Gregory Zeikus, Cloning, Sequencing, and Expression of the Gene Encoding Extracellular a-Amylase from Pyrococcus furiosus and Biochemical Characterization of the Recombinant Enzyme. Appl.Enviro.Microbiol, 1997, 63(9): 3569-3576
42. Hershberger PA, Dickson JA, Friesen PD, Site-specific mutagenesis of the 35K protein gene encoded by Autographa californica Nuclear Polyhedrosis Virus: cell line -specific effects on virus replication, J Viro, 1992, 66:5525-5533
43. Hideo Beniya, Sharon C Braunagel, Max D Summers, Autographa californica nuclear polyhedrosis virus: subcellular localization and protein trafficking of BV/ODV-E26 tointranuclear membranes and viral envelopes, Virol, 1998, 240:64-75
44. Hofmann C, Sandig V, Jennings G, Rudolph M, Schlag P, Strauss M, Efficient gene transfer into human hepatocytes by baculovirus vectors, Proc Natl Acad Sci U S A, 1995, 92: 10099-10103
45. Hu ZH, Broer R, Westerlaken J, Martens JW, Jin F, Jehle JA, Wang LM, VlaK JM, Characterization of the ecdysteroid UDP-glucosyltransferase gene of a single nucleocapsid nucleopolyhedrovirus of Buzura suppressaria, Virus Res, 1997, 47:91-97
46. Ijkel WF, van Strien EA, Heldens JG, Broer R, Zuidema D, Goldbach RW and Vlak JM, Sequence and organization of the Spodoptera exigua multicapsid nucleopolyhedrovirus genome, J Gen Virol, 1999, 80:3289-3304
47. Ingle MB, Erickon R J, Bacterial alpha-amylase, Adv Appl microbial, 1978, 24:257-258
48. Jespersen, H M, E A MacGregor, B Henrissat, M R Sieks, B Svensson, Starch- and glycogen-debranching and branching enzymes: prediction of structural features of the catalytic (β/α)8-barrel domain and evolutionary relationship of other amylolytic enzymes, J Protein Chem, 1993, 12:791-805
49. J kel W F, van Strien E A, Heldens J G, et al., Sequence and organization of the Spodoptera exigua nulticapsid nucleopolyhedrovirus genome, J Gen Virol, 1999, 80:3289 - 3304
50. Johannes TM Flipsen, Ruudm W Mans, Arjen WF Kleefsman, Dangmar knebel- Morsdorf, Just M valk, Deletion of the baculovirus ecdysteroid UDP-glucosyltransferase gene induces early degeneration of malpighian tubules in infected insects, J virol, 1995, 69(7): 4529-4532
51. Kanno M, A Baidocaldarius a-amylase that is highly stable to heat under acidic conditions. Agric Biol Chem, 1986, 50:23-31
52. Kirk B, R Kuchta, Arg304 of human DNA primase is a key contributor to catalysis and NTP binding: primase and the family X polymerases share significant sequence homology, Biochemistry, 1999, 38:7727-7736
53. Koivula TT, Hemila H, Pakkanen R, Sibakov M, Palva 1, Cloning and sequencing of a gene encoding acidophilic amylase from Bacillus acidocaldarius, J Gen Microbiol, 1993, 139(10): 2399-2407
54. Kuzio J, Pearson MN, Harwood SH, Funk C J, Evans JT, Slavicek JM, Rohrmann GF, Sequence and analysis of the genome of a baculovirus pathogenic for Lymantria dispar, Virology, 1999, 253:17-34
55. Li Yonghong, Passarelli AL, Miller L, et al, Identification, Sequence, and Transcriptional Mapping of lef-3, a Baculovirus Gene Involved in Late and Very Late Gene Expression, Journal of Virology, 1993, 67(9): 5260-5268
56. Maeda S, Majima K, Molecular cloning and physical mapping of the genome of Bombyx mori nuclear polyhedrosis virus, J Gen Virol, 1990, 71:1851-1855
57. Martin DA, Stwphen CH, John Kuzio, et al., The Complete DNA Sequence of Autographa californica Nuclear Polyhedrosis Virus, Virology, 1994, 202:586-605
58. Matzke J, Schwermann B, Bakker EP, Acidostable and acidophilic proteins: the example of the alpha-amylase from Alicyclobacillus acidocaldarius, Comp Biochem Physiol A Physiol, 1997, 118(3):,475-479
59. Motoo A, Tatsuo K, Hiroshi O, Yasuji M, Koichi Y, Acid-stable a-amylase of black Aspergilli Part Ⅳ. Some Physicochemical Properties, Agric boil chem, 1968, 32(4): 507-513
60. O'Reilly DR, Brown MR, Miller LK, Alteration of ecdysteroid metabolism due to baculovirus infection of the fall armyworm Spodoptera frugiperda: host ecdysteroid sare conjugated with galactose, Insect Biochem Mol Biol, 1992, 22:313-320
61. O'Reilly DR, Howarth OW, Rees HH, Structure of the ecdysone glucoside formed by a baculovirus ecdysteroid UDP-glucosyltransferase, Insect Biochem, 1991, 21:795-801
62. O'Reilly DR, Miller LK, A baculovirus blocks insect molting by producing ecdysteroid UDP-glucosyltransferase, Science, 1989, 245:1110-1112
63. O'Reilly DR, Miller LK, Regulation of expression of a baculovirus ecdysteroid UDPglucosyltransferase gene, J Virol, 1990, 64(3): 1321-1328
64. O'Reilly DR, Passarelli AL, Goldman IF, Miller LK, Characterization of the DA26 gene in a hypervariable region of the Autographa californica nuclear polyhedrosis virus genome, J Gen Virol, 1990, 71 (5): 1029-1037
65. Passareli AL, Miller LK, Identification and characterization of lef-1, a baculovirus gene involved in late and very late gene expression, J Virol, 1993, 67(6): 3481-3488
66. Pearson MN, Bjornson R, Ahrens C, Rohrmann GF, Identification and characterizeation of a putative origin of DNA replication in the genome of a baculovirus pathogenic for Orgyia pseudotsugata, Virology, 1993, 197(2): 715-725
67. Pennock GD, Shoemaker C, Miller LK, Strong and regulated expression of Escherichia coli beta-galactosidase in insect cells with a baculovirus vector, Mol Cell Biol, 1984, 4(3): 399-406
68. Riegel CL, Lanner-Herrera C, Slavicek JM, Identification and characterization of the ecdysteroid UDP-glucosyltransferase gene of the Lymantria dispar multinucleocapsid nuclear polyhedrosis virus, J Gen Virol, 1994, 75 ( Pt 4): 829-838
69. Robert X, Gottschalk TE, Haser R, et al., Expression, purification and preliminary crystallographic studies of a-amylase isozyme 1 from barley seeds. Acta Crystallogr D Biol Crystallogr, 2002, 58(Pt4): 683-686
70. Sandig V, Hofmann C, Steinert S, Jennings G, Schlag P, Strauss M, Gene transfer into hepatocytes and human liver tissue by baculovirus vectors, Hum Gene Ther. 1996, 7: 1937-1945
71. Schwermann B, Pfau K, Liliensiek B, Schleyer M, Fischer T, Bakker EP, Purification, properties and structural aspects of a thermoacidophilic alpha-amylase from Alicyclobacillus acidocaldarius atcc 27009. Insight into acidostability of proteins, Eur J Biochem, 1994, 226(3): 981-991
72. Shigeaki M, Kimio I, Satoshi S, Toyoaki S, Purification and Some Properties of Acidstable α-amylase from Shochu koji(Aspergillus kawachii), Agric boil chem, 1987, 51(9): 2495-2501
73. Shoji I-Aizaki H, Tani H, lshii K, Chiba T, Saito I, Miyamura T, Matsuura Y, Efficient gene transfer into various mammalian cells, including nonhepatic cells, by baculovirus vectors, J Gen Virol, 1997, 78:2657-2664
74. Smith I, Goodale C, Sequence and in vivo transcription of Lacanobia oleracea granulovirus egt, J Gen Virol, 1998, 79:405-413
75. Smith GE, Summers MD, Fraser M J, Production of human beta interferon in insect cells infected with a baculovirus expression vector, Mol Cell Biol, 1983, 3(12): 2156-2165
76. Theilmann DA, Stewart S, molecular analysis of the trans-acting ie-2 gene of Orgyia pseudotsugata Multinucleocapsid Nuclear Polyhedrosis Virus, Virology, 1992, 187:84-96
77. Thiem SM, miller LK, Identification, Sequence, and Transcriptional Mapping of the major capsid protein gene of the Baculovirus Autographa californica Nuclear Polyh- edrosis Virus, J Viro, 1989, 63: 2008-2018
78. Toister Achitur M, Fakter O, Transcriptional analysis and promoter activity of the Spodoptera littoralis multicapsid nucleopolyhedrovirus ecdysteroid UDP-glucosyltra- nsferase gene, J Gen Virol, 1997, 78:487-491
79. Uchino F, A thermophilic and unusually acidophilic amylase produced by a thermophilic acidophilic Bacillus sp. Agric Biol Chem, 1982, 46(1): 7-13
80. Valk JM, Klinkenberg FA, Zaal K JM, et al., Functional studies on the plO gene of Autographa californica Nuclear Polyhedrosis Virus using a recombinant expressing a p10-beta-galactosidase fusion gene, J Gen Viro, 1988, 69:765-776
81. Victor S.Mikhailov, George F.Rohrmann, Baculovirus replication factor LEF-I is a DNA primase, J Virol, 2002, 76(5): 2287-2297
82. Violet M, J-C Meunier, Kinetic study of the irreversible thermal denaturation of Bacillus licheniformis a-amylase, Biochem J, 1989, 263:665-570
83. Xin Hang, Wen Dong, Linda A, Guarino, The lef-3 Gene of Autographa californiea Nuclear Polyhedrosis Virus Encodes a Single-Stranded DNA-binding Protein, Journal of Virology, 1995, 69(6): 3924-2928
84. Yasuji M, Motoo A, Yasushi T, Koichi Y, Acid-stabe a-amylase of Black Aspergilli Part Ⅲ. Separation of Acid-stabe a-amylase and Acid-unstabe a-amylase from the Same Mold Amylasepreparation, Agric boil chem, 1968,32(1):110-113
85. Yasuji M, Tatsuo K, Motoo A, Koichi Y, Acid-stable a-amylase of Black Aspergilli Part Ⅱ. Some General Properties, Agric boil chem, 1968, 32(1): 104-109
2.储瑞银,必怡德,吕鸿声,李载平,吴祥甫,乙型肝炎病毒表面抗原基因(pres2-S)在家蚕培养细胞中的表达,生物化学和生物物理学报,1990,22:385-389
3.洪华珠,杨红,中国生物防治,1995,11(2):84-88
4.李镇,龙綮新,袁美妗,贺雄雷,庞义,王珣章,斜纹夜蛾核多角体病毒(SpltNPV)基因组EcoR I-G片段的序列分析,中国病毒学,2002,17(2):119-124
5.林建城,酶在食品工业、轻工业和环境保护上的应用分析,莆田学院学报,2005,12(2):17-20
6.沈兴家,BmNPV和AcNPV egt基因启动子特性及家蚕海藻糖基因启动子活性的滞育激素调节研究[博士学位论文],北京:中国农业科学院研究生院,2004
7.石生林,柞蚕核型多角体病毒B-ORF6L、ptp、lef-12基因克隆及表达[博士学位论文],北京:中国科学院,2005
8.王海燕,秦浚川,王敖全,唐国敏,黑曲霉酸性α-淀粉酶基因和糖化酶基因对工业酒精酵母的整合及其共表达,微生物学报,2004,44(4):483-486
9.王文兵,张志芳,何家禄,吕鸿声,杆状病毒基冈组DNA复制基因的研究进展,生物化学与生物物理进展,2000,27(3):257-261
10.袁铁铮,姚斌,罗会颖,王亚茹,伍宁丰,范云六,来源于嗜热脂环酸杆菌的嗜酸性α-淀粉酶的研究,生物工程学报,2005,21(1):78-83
11.张春发,刘淑珊,范琦,李广泽,柞蚕核型多角体病毒载体表达系统基因工程研究进展,华南农业大学学报,1995,16(1):133-137
12.张纪忠,微生物分类学,上海:复旦大学出版社,1990,377-412
13.张丽苹,徐岩,金建中,酸性α-淀粉酶的研究与应用,酿酒,2002,29(3):19—21
14.张小霞,梁振普,牛国栋,等,棉铃虫核多角体病毒lef-3基因的序列分析,中国病毒学,2002,17(3):234—238
15.张志芳,何家禄,吴祥甫,家蚕核多角体病毒DNA聚合酶基因的克隆,蚕业科学,1995,21(2):86-90
16.张志芳,张颖,吕鸿声,等,家蚕核多角体病毒解链酶基因的克隆及部分序列分析,病毒学报,1994,10(4):381-383
17.张志芳,张颖,吕鸿声等,家蚕核多角体病毒DNA复制起始点hr3的结构功,中国科学B辑,1995,25(9):949-955
18. Ahrens CH, Rohrmann GF, Replication of Orgyia pseudotsugata baculovirus DNA: lef-2 and ie-1 are essential and ie-2, p34, and Op-iap are stimulatory genes, Virology, 1995, 212(2): 650-662
19. Ahrens CH, Russell R LQ, Funk CJ, The Sequence of the Orgyia pseudotsugata Multinucleocapsid Nuclear Polyhedrosis Virus Genome, Virology, 1997, 229:381-399
20. Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ, Basic local alignment search tool, J Mol Biol, 1990, 215:403-410
21. Antje C, Springer Handbook of Enzymes, 2nded, New York: Springer-Verlag Berlin Heidelberg, 2003, 1-39
22. Augustin M, R Huber, J Kaiser, Crystal structure of a DNA dependent RNA polymerase(DNA primase), Nat Struct Biol, 2001, 8:57-61
23. Barrett JW, Krell PJ, Arif BM, Characterization, sequencing and phylogeny of the ecdysteroid UDP-glucosyltransferase gene from two distinct nuclear polyhedrosis viruses isolated from Choristoneurafumiferana, J Gen Virol, 1995, 76:2447-2456
24. Blissard GW, Quant-Russell RL, Rohrmann GF, Beaudreau GS, Nucleotide Sequence, Transcriptional Mapping, and temporal expression of the gene encoding p39, a major structural protein of the muticapisid Nuclear Polyhedrosis Virus of Orgyia pseudotsugata, Virology, 1989, 168:354-362
25. Boyce FM, Bucher NL, Baculovirus-mediated gene transfer into mammalian cells, Proc Natl Acad Sci U S A, 1996, 93:2348-2352
26. Buonocore V, Caporale C, De Rosa M, Gambacorta A, Stable, inducible thermoacidophilic alpha-amylase from Bacillus acidocaldarius, J Bacteriol, 1976, 128(2):515-21
27. Carstens EB, Lu AL, Chan HB, Sequence, transcriptional mapping, and over expression of p47, a baculovirus gene regulating late gene expression, J Viro, 1993, 67:2513-2520
28. Chert XW, IJkel WF, Tarchini R, XL Sun, Sandbrink H, H Wang, Peters S, Zuidema D, Lankhorst RK, Vlak JM and ZH H, The sequence of the Helicoverpa armigera single nucleocapsid nucleopolyhedrovirus genome, J Gen Virol, 2001, 82:241-257
29. Christian HA, Cheryl Carlson, George FR, Identification, Sequence, and Transcriptional Analysis of lef-3, a Gene Essential for Orgyia pseudotsugata Baculovirus DNA Replication(J). Virology, 1995,210:372-382
30. Clarke EE, Tristem M, Cory JS, O'Reilly DR, Characterization of the ecdysteroid UDP-glucosyltransferase gene from Mamestra brassicae nucleopolyhedrovirus, J Gen Virol, 1996, 77:2865-2871
31. Clem RJ, Fechheimer M, Miller LK, Prevention of apoptosis by a baculovirus gene during infection of insect cells, Science, 1991,254:1388-1390
32. Copeland W, X Tan, Active site mapping of the catalytic mouse primase subunit by alanine scanning mutagenesis, J Biol Chem, 1995,270:3905-3913
33. Christian H A,Cheryl Carlson, George F R, Identification, Sequence, and Trans- criptional Analysis of lef-3, a Gene Essential for Orgyia pseudotsugata Baculovirus DNA Replication, Virology, 1995, 210:372-382
34. Copeland W, X Tan, Active site mapping of the catalytic mouse primase subunit by alanine scanning mutagenesis, J Biol Chem, 1995, 270:3905-3913
35. Dingwall C, Laskey RA, Nuclear targeting sequences-A consensus? TIBS 1991, 16:478-481
36. Evans JT, Leisy D J, Rohrmann GF, Characterization of the interaction between the baculovirus replication factors LEF-1 and LEF-2, J Virol, 1997, 71(4): 3114-3119
37. Flipsen JTM, Deletion of the baculovirus ecdysteroid UDP-glucosyltransferase gene induces early degeneration of Malpighian tubules on infected insects, J Virol, 1995, 69:4529-4532
38. Friesen PD, Miller LK, The regulation of baculovirus gene expression, Curr Top Microbiol Immunol, 1986, 131:31-49
39. Fukusumi, S, A Kamizono, S Horinouchi, T Beppu, Cloning and nucleotide sequence of a heat-stable amylase gene from an anaerobic thermophile, Dictyoglomus thermophilum, Eur J Biochem, 1988, 174:15-21
40. Gomi S, Majima K, Maeda S, Sequence analysis of the genome of Bombyx mori nucleopolyhedrovirus, J Gen Virol, 1999, 80(5): 1323-1337
41. Guoqlang Dong, Clatre Vieille, Alexei Savchenk, J Gregory Zeikus, Cloning, Sequencing, and Expression of the Gene Encoding Extracellular a-Amylase from Pyrococcus furiosus and Biochemical Characterization of the Recombinant Enzyme. Appl.Enviro.Microbiol, 1997, 63(9): 3569-3576
42. Hershberger PA, Dickson JA, Friesen PD, Site-specific mutagenesis of the 35K protein gene encoded by Autographa californica Nuclear Polyhedrosis Virus: cell line -specific effects on virus replication, J Viro, 1992, 66:5525-5533
43. Hideo Beniya, Sharon C Braunagel, Max D Summers, Autographa californica nuclear polyhedrosis virus: subcellular localization and protein trafficking of BV/ODV-E26 tointranuclear membranes and viral envelopes, Virol, 1998, 240:64-75
44. Hofmann C, Sandig V, Jennings G, Rudolph M, Schlag P, Strauss M, Efficient gene transfer into human hepatocytes by baculovirus vectors, Proc Natl Acad Sci U S A, 1995, 92: 10099-10103
45. Hu ZH, Broer R, Westerlaken J, Martens JW, Jin F, Jehle JA, Wang LM, VlaK JM, Characterization of the ecdysteroid UDP-glucosyltransferase gene of a single nucleocapsid nucleopolyhedrovirus of Buzura suppressaria, Virus Res, 1997, 47:91-97
46. Ijkel WF, van Strien EA, Heldens JG, Broer R, Zuidema D, Goldbach RW and Vlak JM, Sequence and organization of the Spodoptera exigua multicapsid nucleopolyhedrovirus genome, J Gen Virol, 1999, 80:3289-3304
47. Ingle MB, Erickon R J, Bacterial alpha-amylase, Adv Appl microbial, 1978, 24:257-258
48. Jespersen, H M, E A MacGregor, B Henrissat, M R Sieks, B Svensson, Starch- and glycogen-debranching and branching enzymes: prediction of structural features of the catalytic (β/α)8-barrel domain and evolutionary relationship of other amylolytic enzymes, J Protein Chem, 1993, 12:791-805
49. J kel W F, van Strien E A, Heldens J G, et al., Sequence and organization of the Spodoptera exigua nulticapsid nucleopolyhedrovirus genome, J Gen Virol, 1999, 80:3289 - 3304
50. Johannes TM Flipsen, Ruudm W Mans, Arjen WF Kleefsman, Dangmar knebel- Morsdorf, Just M valk, Deletion of the baculovirus ecdysteroid UDP-glucosyltransferase gene induces early degeneration of malpighian tubules in infected insects, J virol, 1995, 69(7): 4529-4532
51. Kanno M, A Baidocaldarius a-amylase that is highly stable to heat under acidic conditions. Agric Biol Chem, 1986, 50:23-31
52. Kirk B, R Kuchta, Arg304 of human DNA primase is a key contributor to catalysis and NTP binding: primase and the family X polymerases share significant sequence homology, Biochemistry, 1999, 38:7727-7736
53. Koivula TT, Hemila H, Pakkanen R, Sibakov M, Palva 1, Cloning and sequencing of a gene encoding acidophilic amylase from Bacillus acidocaldarius, J Gen Microbiol, 1993, 139(10): 2399-2407
54. Kuzio J, Pearson MN, Harwood SH, Funk C J, Evans JT, Slavicek JM, Rohrmann GF, Sequence and analysis of the genome of a baculovirus pathogenic for Lymantria dispar, Virology, 1999, 253:17-34
55. Li Yonghong, Passarelli AL, Miller L, et al, Identification, Sequence, and Transcriptional Mapping of lef-3, a Baculovirus Gene Involved in Late and Very Late Gene Expression, Journal of Virology, 1993, 67(9): 5260-5268
56. Maeda S, Majima K, Molecular cloning and physical mapping of the genome of Bombyx mori nuclear polyhedrosis virus, J Gen Virol, 1990, 71:1851-1855
57. Martin DA, Stwphen CH, John Kuzio, et al., The Complete DNA Sequence of Autographa californica Nuclear Polyhedrosis Virus, Virology, 1994, 202:586-605
58. Matzke J, Schwermann B, Bakker EP, Acidostable and acidophilic proteins: the example of the alpha-amylase from Alicyclobacillus acidocaldarius, Comp Biochem Physiol A Physiol, 1997, 118(3):,475-479
59. Motoo A, Tatsuo K, Hiroshi O, Yasuji M, Koichi Y, Acid-stable a-amylase of black Aspergilli Part Ⅳ. Some Physicochemical Properties, Agric boil chem, 1968, 32(4): 507-513
60. O'Reilly DR, Brown MR, Miller LK, Alteration of ecdysteroid metabolism due to baculovirus infection of the fall armyworm Spodoptera frugiperda: host ecdysteroid sare conjugated with galactose, Insect Biochem Mol Biol, 1992, 22:313-320
61. O'Reilly DR, Howarth OW, Rees HH, Structure of the ecdysone glucoside formed by a baculovirus ecdysteroid UDP-glucosyltransferase, Insect Biochem, 1991, 21:795-801
62. O'Reilly DR, Miller LK, A baculovirus blocks insect molting by producing ecdysteroid UDP-glucosyltransferase, Science, 1989, 245:1110-1112
63. O'Reilly DR, Miller LK, Regulation of expression of a baculovirus ecdysteroid UDPglucosyltransferase gene, J Virol, 1990, 64(3): 1321-1328
64. O'Reilly DR, Passarelli AL, Goldman IF, Miller LK, Characterization of the DA26 gene in a hypervariable region of the Autographa californica nuclear polyhedrosis virus genome, J Gen Virol, 1990, 71 (5): 1029-1037
65. Passareli AL, Miller LK, Identification and characterization of lef-1, a baculovirus gene involved in late and very late gene expression, J Virol, 1993, 67(6): 3481-3488
66. Pearson MN, Bjornson R, Ahrens C, Rohrmann GF, Identification and characterizeation of a putative origin of DNA replication in the genome of a baculovirus pathogenic for Orgyia pseudotsugata, Virology, 1993, 197(2): 715-725
67. Pennock GD, Shoemaker C, Miller LK, Strong and regulated expression of Escherichia coli beta-galactosidase in insect cells with a baculovirus vector, Mol Cell Biol, 1984, 4(3): 399-406
68. Riegel CL, Lanner-Herrera C, Slavicek JM, Identification and characterization of the ecdysteroid UDP-glucosyltransferase gene of the Lymantria dispar multinucleocapsid nuclear polyhedrosis virus, J Gen Virol, 1994, 75 ( Pt 4): 829-838
69. Robert X, Gottschalk TE, Haser R, et al., Expression, purification and preliminary crystallographic studies of a-amylase isozyme 1 from barley seeds. Acta Crystallogr D Biol Crystallogr, 2002, 58(Pt4): 683-686
70. Sandig V, Hofmann C, Steinert S, Jennings G, Schlag P, Strauss M, Gene transfer into hepatocytes and human liver tissue by baculovirus vectors, Hum Gene Ther. 1996, 7: 1937-1945
71. Schwermann B, Pfau K, Liliensiek B, Schleyer M, Fischer T, Bakker EP, Purification, properties and structural aspects of a thermoacidophilic alpha-amylase from Alicyclobacillus acidocaldarius atcc 27009. Insight into acidostability of proteins, Eur J Biochem, 1994, 226(3): 981-991
72. Shigeaki M, Kimio I, Satoshi S, Toyoaki S, Purification and Some Properties of Acidstable α-amylase from Shochu koji(Aspergillus kawachii), Agric boil chem, 1987, 51(9): 2495-2501
73. Shoji I-Aizaki H, Tani H, lshii K, Chiba T, Saito I, Miyamura T, Matsuura Y, Efficient gene transfer into various mammalian cells, including nonhepatic cells, by baculovirus vectors, J Gen Virol, 1997, 78:2657-2664
74. Smith I, Goodale C, Sequence and in vivo transcription of Lacanobia oleracea granulovirus egt, J Gen Virol, 1998, 79:405-413
75. Smith GE, Summers MD, Fraser M J, Production of human beta interferon in insect cells infected with a baculovirus expression vector, Mol Cell Biol, 1983, 3(12): 2156-2165
76. Theilmann DA, Stewart S, molecular analysis of the trans-acting ie-2 gene of Orgyia pseudotsugata Multinucleocapsid Nuclear Polyhedrosis Virus, Virology, 1992, 187:84-96
77. Thiem SM, miller LK, Identification, Sequence, and Transcriptional Mapping of the major capsid protein gene of the Baculovirus Autographa californica Nuclear Polyh- edrosis Virus, J Viro, 1989, 63: 2008-2018
78. Toister Achitur M, Fakter O, Transcriptional analysis and promoter activity of the Spodoptera littoralis multicapsid nucleopolyhedrovirus ecdysteroid UDP-glucosyltra- nsferase gene, J Gen Virol, 1997, 78:487-491
79. Uchino F, A thermophilic and unusually acidophilic amylase produced by a thermophilic acidophilic Bacillus sp. Agric Biol Chem, 1982, 46(1): 7-13
80. Valk JM, Klinkenberg FA, Zaal K JM, et al., Functional studies on the plO gene of Autographa californica Nuclear Polyhedrosis Virus using a recombinant expressing a p10-beta-galactosidase fusion gene, J Gen Viro, 1988, 69:765-776
81. Victor S.Mikhailov, George F.Rohrmann, Baculovirus replication factor LEF-I is a DNA primase, J Virol, 2002, 76(5): 2287-2297
82. Violet M, J-C Meunier, Kinetic study of the irreversible thermal denaturation of Bacillus licheniformis a-amylase, Biochem J, 1989, 263:665-570
83. Xin Hang, Wen Dong, Linda A, Guarino, The lef-3 Gene of Autographa californiea Nuclear Polyhedrosis Virus Encodes a Single-Stranded DNA-binding Protein, Journal of Virology, 1995, 69(6): 3924-2928
84. Yasuji M, Motoo A, Yasushi T, Koichi Y, Acid-stabe a-amylase of Black Aspergilli Part Ⅲ. Separation of Acid-stabe a-amylase and Acid-unstabe a-amylase from the Same Mold Amylasepreparation, Agric boil chem, 1968,32(1):110-113
85. Yasuji M, Tatsuo K, Motoo A, Koichi Y, Acid-stable a-amylase of Black Aspergilli Part Ⅱ. Some General Properties, Agric boil chem, 1968, 32(1): 104-109