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1 HC-EGFP 200 HC-EGFPBombyx morihc-egfp i
2 piggybac ii
3 iii
4
5 HC-EGFPBombyx mori HC-EGFP HC-EGFP (1) 4 m 1.5 m (2) (3) 4 mm (4) (5) (1) 2 (2) (3) 2
6 5 silkworm Bombyx mori (Linnaeus) 1 DNA w-1 pnd 15 2 /C w-1 pnd /C
7 w-1 pnd w-1 pnd 5 DNA w-1 / + w-1 w-1 / w-1 3 4
8 w-1 pnd pnd No.848 w-1 No /C w-1 pnd /C A A pnd pnd + + A + pnd A A A A A pnd pnd pnd A A A /C 5
9 /C 5 pndw-1w-1 pnd + pnd + w-1 CS01 + pnd w w-1 pnd /C CK01 CT GNK2 GNT3 25 Bombyx mandarina 1 30 w-1 pnd /C
10 B. mori B. mandarina ; 1995; 1998; ; 2013;
11 80 77%62 16% ; kg12, , ,029 6,
12 Nosema bombycis
13 mm1 mm0.5 mm cm 85%
14 %50% m cm , kg m 3.5 m
15
16 ; , % , %
17 F 3 F ; 1939 F Kuwahara et al., % 89% % %20 85% %20 9 0%
18 /C w-1 pnd3 Blepharipa zebinaexorista sorbillanspediculoides ventricosus 1992 Exorista japonica piggybac 20 piggybac Trichoplusia ni TN-368 DNA Cary et al., 1989; Handler, 2002 piggybac 2 piggybac piggybac piggybac piggybac piggybac piggybac 1 15
19 piggybac 12piggyBac
20 1 5 HC-EGFPBombyx morihc-egfp HC-EGFP H HC-EGFP Fibroin H promoter1.1 kb H H H HC-EGFP Kojima et al., 2007 HC-EGFP2.3 kb H H Aequorea victoria pegfp EGFP Kojima et al., 2007 Fibroin H polya0.3 kb H H Kojima et al., 2007 AB xP3 promoter 0.2 kb 3xP3 hsp70 DsRed2 17
21 Berghammer et al., 1999; Thomas et al., 2002 DsRed20.7 kb Discosoma sp. ; Matz et al., xP3 promoter DsRed2 SV40 polyasv kb 40Simian virus 40 AB piggybac R 1.1 kb Trichoplusia ni piggybac Cary et al., 1989 piggybac piggybac L 0.7 kb Trichoplusia ni piggybac Cary et al., 1989 piggybac puc ori 0.7 kb ColE1 AmpR 0.9 kb 18
22 5 15 piggybac R 14HC-EGFP pbac[3xp3-dsred2afm]_hc-egfp piggybac piggybac 3xP3 DsRed2 SV40 polya HC-EGFP 3xP3 40 Fibroin H promoter H HC-EGFP Fibroin H polya piggybac L H EGFP H piggybac piggybac puc ori AmpR RT-PCR RT-PCR 15 piggybac puc18 p3e1.2cary et al, 19893xP3-DsRed2 20 piggybac pbac[3xp3-dsred2afm] Inoue et al, 2005 H HC-EGFP pbac[3xp3-dsred2afm]_hc-egfp Kojima et al,
23 Fibroin H promoter Fibroin H Fibroin H polya H EGFP HC-EGFP HC-EGFP pbac[3xp3-dsred2afm]_hc-egfp pbac[3xp3-dsred2afm] piggybac AmpR puc18 puc ori 3xP3 promoter DsRed2 p3e1.2 SV40 plya 15HC-EGFP pbac[3xp3-dsred2afm]_hc-egfp 5 piggybacr piggybacl 13 pha3pig pha3pig 2 16 Tamura et al., pHA3PIG A3 promoter0.7 kb A3 piggybac transposasepiggybac Mounier and Prudhomme, kb Trichoplusia ni piggybac Cary et al., 1989 piggybac 2 DNA 20
24 piggybac R 1.1 kb Trichoplusia ni piggybac Cary et al., 1989 piggybac AmpR 0.9 kb puc ori 0.7 kb ColE1 16pHA3PIG A3 promoter A3 5 piggybac transposase piggybac piggybac R piggybac piggybac puc ori AmpR PCR PCR HC-EGFP 15 HC-EGFP H Aequorea victoria 21
25 H H mrna Takiya et al., 1990; Kojima et al., 2007 Green Fluorescent ProteinGFP 5 EGFPEnhanced GFP HC-EGFP H EGFP Kojima et al., 2007 EGFP Food Allergy Research and Resource Program DatabaseFARRPver FASTA 15 DsRed2 DsRed2 Discosoma sp. 20 3xP3 3xP3 Drosophila melanogaster hsp70 3xP3 Sheng et al., 1997; Thomas et al., xP3 25 SV40 40 mrna DsRed2 Food Allergy Research and Resource Program DatabaseFARRPver FASTA
26 5 A3 piggybac piggybac 1 13 Escherichia coli puc18 piggybac puc18 p3e1.2 Cary et al, 1989; 15 pbac[3xp3-dsred2afm]_hc-egfp p3e1.2 HC-EGFP 15 2 piggybac pha3pig Tamura et al., 2000; 16 pha3pig A3 15 piggybac 20 puc18 2,686 bpl08752 puc18 piggybacp3e1.2 5,958 bp piggybac Website pbac[3xp3-dsred2afm]_hc-egfp,306 bp 8 25 pha3pig 6,160 bp9 puc18 30 p3e1.2 piggybac piggybac pbac[3xp3-dsred2afm]_hc-egfp p3e1.2 piggybac 35 pha3pig 23
27 A3 piggybac A3 piggybac piggybac pbac[3xp3-dsred2afm]_hc-egfp piggybac DsRed2 HC-EGFP pbac[3xp3-dsred2afm]_hc-egfp14pha3pig 1617 piggybac 2 1 piggybac
28 17 5 pbac[3xp3-dsred2afm]_hc-egfp pha3pig G G 0 /C G
29 /C w1 pnd G0 G1 /C 200 G2-2 G1-2 /C 200 G3-3 G G3-4 G3-5 G2-4 G1-3 G3-6 G2-5 G3-7 HC-EGFP 200 G3-8 G2-6 G3-11 G2-9 HC-EGFP G1-4 G1-8 KH25 G3-12 G4-1 G2- G1-9 G3-13 G4-2 G2-11 G HC-EGFP 200 HC-EGFP 1 26
30 3 18 G 1-6 G 1-8 G 2- G 3-12 G 4-1 G 4-2 G PCR RT-PCR piggybac 5 pbac[3xp3-dsred2afm]_hc-egfp piggybac pbac[3xp3-dsred2afm]_hc-egfp piggybac AcNPVAutographa california nucleopolyhedrovirus FP 642 bp5' 340 bplef9 1,548 bp5'
31 bp piggybac 5 G DNA PCR PCR 16 DNA piggybac P1ADsRed2 EGFP 200HC-EGFP HC-EGFP P1A /CKH KH25/CF 1 /C 1: G 1 5 DNA piggybac 35 28
32 G 1 5 RNA EGFP RT-PCR 5 13 EGFP g DNA 15 HC-EGFP H HC-EGFP H 20 3xP3 DsRed2 HC-EGFP H EGFP 25 DsRed2 30 HC-EGFP HC-EGFP 200HC-EGFP HC-EGFP
33 HC-EGFP 200 HC-EGFP 200 HC-EGFP HC-EGFP P g 2.17 g0.271 g0.551 g %98.1% 16P = %98.8% 2 18 P =
34 20 cm cm5.4 cm 9.6 cm5.0 cm 5 19P = cm P < cm 15 21P < cm 93% 18 cm 85% EGFP DsRed
35 EGFP DsRed EGFP DsRed2 GFP 15 Toxin and Toxin Target Database Food Allergy Research and Resource Program DatabaseFARRP ver HC-EGFP HC-EGFP
36 (1) 15 4 m1.5 m (2) (3) 20 4 mm (4) (5) (1) 2 (2) 30 (3)
37 5 34
38 ; H HC-EGFP % 35 35
39 H HC-EGFP
40 5 H HC-EGFP EGFP DsRed Bombyx huttoni 37
41 Hutton, 1864; 1998; 1999; ; 1939; 7 Kuwahara et al., 1984; Daimon et al., ; ; 1997;
42 A. 15,
43 15, B C D E. 1 20,000 5, ,000 AD
44 cox1cytochrome c oxidase I4, Yukuhiro et al., 2012a ; 1997;
45 Bombyx mori 5 H HC-EGFP H HC-EGFP H HC-EGFP 42
46
47 H HC-EGFP
48 Berghammer A. J., Klingler M., and Wimmer E. A. (1999) A universal marker for transgenic insects. Nature, 402, Cary L. C., Goebel M., Corsaro B. G., Wang H.-G., Rosen E., and Fraser M. J. (1989) Transposon mutagenesis of baculoviruses: analysis of Trichoplusia ni transposon IFP2 insertions within the FP-locus of nuclear polyhedrosis viruses. Virology, 172, Daimon T., Fujii T., Fujii T., Yokoyama T., Katsuma S., Shinoda T., Shimada T. and Ishikawa Y. (2012) Reinvestigation of the sex pheromone of the wild silkmoth Bombyx mandarina: the effects of bombykal and bombykol acetate. J. Chem. Ecol., 38, Handler A. M. (2002) Use of the piggybac transposon for germ-line transformation of insects. Insect Biochem. Mol. Biol., 32, Hutton T. (1864) On the revision and restoration of the silkworm (part II.); with distinctive characters of eighteen species of silk-producing Bombycidae. Trans. Entmol. Soc. London, 3rd ser., Vol. II, Inoue S., Kanda T., Imamura M., Quan G.-X., Kojima K., Tanaka H., Tomita M., Hino R., Yoshizato K., Mizuno S., and Tamura T. (2005) A fibroin secretion-deficient silkworm mutant, Nd-s D, provides an efficient system for producing recombinant proteins. Insect Biochem. Mol. Biol. 35, Kojima K., Kuwana Y., Sezutsu H., Kobayashi I., Uchino K., Tamura T., and Tamada Y. (2007) A new method for the modification of fibroin heavy chain in the transgenic silkworm. Biosci. Biotechnol. Biochem, 71, Kuwahara Y., Mori N., Yamada S., and Nemoto T. (1984) Evaluation of bombykol as the sex pheromone of Bombyx mandarina (Lepidoptera: Bombycidae). Appl. Entomol. Zool. 19, Matz M. V., Fradkov A. F., Labas Y. A., Savitsky A. P., Zaraisky A. G., Markelov M. L., and Lukyanov S. A. (1999) Fluorescent proteins from nonbioluminescent Anthozoa species. Nature Biotechnol., 17, Mounier N. and Prudhomme J. C. (1991) Differentianl expression of muscle and cytoplasmic actin genes during development of Bombyx mori. Insect Biochem., 21, Sheng G., Thouvenot E., Schmucker D., Wilson D. S. and Desplan C. (1997) Direct regulation of rhodopsin 1 by Pax-6/eyeless in Drosophila: evidence for a conserved function in photoreceptors. Genes Dev., 11, Takiya S., Hui C.-c., and Suzuki Y. (1990) A contribution of the core-promoter and its surrounding regions to the preferential transcription of the fibroin gene in posterior silk gland extracts. EMBO J. 9, Tamura T., Thibert C., Royer C., Kanda T., Abraham E., Kamba M., Kômoto N., Thomas J.-L., Mauchamp B., Chavancy G., Shirk P., Fraser M., Prudhomme, J.-C., and Couble P. (2000) Germline transformation of the silkworm Bombyx mori L. using a piggybac transposon-derived vector. Nature Biotechnol., 18, Thomas J.-L., Da Rocha M., Besse A., Mauchamp B., and Chavancy G. (2002) 3xP3-EGFP marker facilitates screening for transgenic silkworm Bombyx mori L. from the embryonic stage onwards. Insect Biochem. Mol. Biol. 32, Yukuhiro K., Iwata K., Kômoto N., Tomita S., Itoh M., and Kiuchi M. (2012a) Nucleotide sequences of mitochondrial cytochrome C oxidase subunit I (COI) gene show clear differences between the domesticated silkmoth Bombyx mori and the wild mulberry silkmoth Bombyx mandarina from Japan. J. 45
49 Insect Biotechnol. Sericol., 81, , , , (A) (2) , , , , , (3)Wild Silkworm News , , ,
50 * 15 * 20 Bombyx mandarina m
51 PCR EGFP DsRed2 DsRed2 EGFP DsRed2 PCR
52 HC-EGFP, Bombyx mori L. HC-EGFP HC-EGFP
53 m 6 12 DsRed2 EGFP DsRed2 PCR DsRed
54
55
56 5 HC-EGFPBombyx mori HC-EGFP HC-EGFP ,000 5, mm
57
58 ,0 m m 5.5 m79 m m 56 4 mm 7 4 mm 36 m 2 12 m 3 m36 m m1.5 m m 1.6 m12 m 2 1 m m 1.8 m4.9 m m 3.3 m2.5 m m3.5 m0.8 m m 55
59 1 km 5 11 km km 56
60 500 m
61 ha 53.7 m 39.5 m 3 58
62 m 5.5 m 6 59
63
64 5 61
65 (1) HC-EGFP Bombyx morihc-egfp HC-EGFP 200 (2) (3) (4) 3 (5) 1 2 (6) 1 2(3) (7) 1 2(3) (8) (7) (9) (7) () 2 (7) 62
66 5 (11) (7) (1) 30 4 mm 6 15 (2) (3) (4) (5) (6) (7) 6 15 (8) 1 (7) (9) 30 63
67
68 1 5 Bombyx mori (Linnaeus) Bombyx mandarina (Moore) 1998; 1999; ; ;
69 ; 1939b Bombyx Bombyx mandarina ; Yukuhiro et al., n = 562n = ; 1999; ; m280m ,000m
70 Nosema bombycis % % n = 56 2n = ; Nakamura et al., 1999; Kawanishi et al., mm1 mm 1936; 1950; ; ; ; ; ; a; Sasaki and Jibiki,
71 ; ; ; Kuwahara, 1984 Kuwahara et al., 1984; Daimon et al., Sasaki and Jibiki, 1984; Kuwahara, ; 1939b ; a; %
72 % Daimon T., Fujii T., Fujii T., Yokoyama T., Katsuma S., Shinoda T., Shimada T. and Ishikawa Y. (2012) Reinvestigation of the sex pheromone of the wild silkmoth Bombyx mandarina: the effects of bombykal and bombykol acetate. J. Chem. Ecol., 38, Kawanishi Y., Banno Y., Fujimoto H., Nho S. K., Tu Z., Mita K., Tsuchida K., Takada N., Maekawa H., and Nakajima Y. (2008) Method for rapid distinction of Bombyx mandarina (Japan) from B. mandarina (China) based on rdna sequence differences. J. Insect Biotechnol. Sericol. 77, Kuwahara Y. (1984) Flight time of Bombyx mandarina males to a pheromone trap baited with bombykol. Appl. Entomol. Zool., 19, Kuwahara Y., Mori N., Yamada S., and Nemoto T. (1984) Evaluation of bombykol as the sex pheromone of Bombyx mandarina (Lepidoptera: Bombycidae). Appl. Entomol. Zool. 19, Nakamura T., Banno Y., Nakada T., Nho S. K., Xü M. K., Ueda K., Kawarabata T., Kawaguchi Y., and Koga K. (1999) Geographic dimorphism of the wild silkworm, Bombyx mandarina, in the chromosome number and the occurrence of a retroposon-like insertion in the arylphorin gene. Genome, 42, Sasaki M. and Jibiki F. (1984) Timing of the sexual behavior of wild and domestic silk moths. Appl. Entomol. Zool., 20, Yukuhiro K., Sezutsu H., Itoh M., Shimizu K., and Banno Y. (2002) Significant levels of sequence divergence and gene rearrangement have occurred between the mitochondrial genome of the wild mulberry silkmoth, Bombyx mandarina, and its close relative, the domesticated silkmoth, Bombyx mori. Mol. Biol. Evol., 19, a 396, b397, , (A) (2) , , , ,
73 195814, (3)Wild Silkworm News ,
74
75 m 20 m 5 1,000 m
76
77 4 1 m 30 cm 5 C8 1 1 (1) (2) % mm 5/ / / / / / / / / / /
78 5/ / / / / / / / / / (3) % mm 7/ / / / / / / / /2 22, / /
79 8/ / / / / / / / / / / / / / / / / (4) % mm 5/ / / (5)
80 % mm 7/ / / / / (6) % mm 9/ / / / / (7) % mm 6/
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84 AB A3 3.2 m 5.2 m 7.6 m m B
85 F F , 21, , 394,
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88 5 G KH25 5 DNA 0.5 g KOD-plus 30 PCR PCR PCR 50 l 5 l % pig-tp4868u25: 5 -TATATCCCAAACAAGCCAAGTAAGT-3 pig-tp5394l23: 5 -CCACCTATTCGTCTTCCTACTGC-3 1 piggybac g 6 pg PCR 550bp0.4 pg DNA 0.5 g M ,000 2,000 1, M 1 DNA 400 ng (DNA 0.5 μg 7x 4 2 DNA 4 ng (700 3 DNA 40 pg (7 4 DNA 0.4 pg ( KH25 DNA 0.5 μg 6 KH25 DNA 0.5 μg5 6 85
89 111 KH25/CF 1 5 /C DNA DNA2 gsali 0.8% Hybond-N+14 piggbac R PCR 15 Rarm-5: 5 -TGTTTTATCGGTCTGTATATCGAGG-3 Rarm-3: 5 -GGTGGCCTATGGCATTATTGTACGG-3 PCR AlkPhos Direct KitGE CDP-Star pbac[3xp3-dsred2afm]6.5 kb AscI (kb)
90
91 12 5 G DNA 11 KH25 (G 1-8 ) 200 HC-EGFP200 (G 3-12 ) KH25 (G 1-8 ) HC-EGFP (G 2- ) (kb) (kb) KH25 (G 1-8 ) x 200 HC-EGFP x HC-EGFP200 (G 4-1 ) (kb)
92 13 G RNA EGFP RT-PCR RNA 1 g AMV Reverse Transcriptase XLcDNA 20 1 Ex Taq HSPCR 14 RT-PCR ks35: 5 -ACGACGGCAACTACAAGACC-3 ks248: 5 -GAACTCCAGCAGGACCATGTGAT-3 PCR l 3 l 2% 15 RT-PCR rp49 NM_ PCR rp49 20 ks13: 5 -GGATCGCTATGACAAACTTAAGAGGA-3 ks12: 5 -TGCTGGGCTCTTTCCACGA-3 365bpEGFP PCR 25 rp bp 89
93 200 EGFP HC-EGFP200 (G 3-12 ) HC-EGFP (G 2- ) EGFP (bp) (bp) rp49 rp49 (bp) (bp) x 200 EGFP HC-EGFP x HC-EGFP200 (G 4-1 ) (bp) rp49 (bp)
94 G 4-2 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP ± ± ± ± ±95.0 x ± ± ± ± ±2 20 mg t P
95 G 4-1 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP 200 g cg % x
96 G 4-1 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP 200 % 1, , x 200 1, , P =
97 G 4-1 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP x
98 G 4-1 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP % x P =
99 cm 20 cm G 4-2 HC-EGFP HC-EGFP cm5.4 cm 9.6 cm5.0 cmt P = cm HC-EGFPHC-EGFP cm 96
100 cm G 4-2 HC-EGFP HC-EGFP HC-EGFP x HC-EGFP 200 x t P <
101 cm G 4-2 HC-EGFP HC-EGFP cm 15 Mann-Whitney U P < HC-EGFP HC-EGFP
102 22 5 1% cm 0 g HC-EGFP HC-EGFP 200 HC-EGFP HC-EGFP % t P HC-EGFP HC-EGFP HC-EGFP HC-EGFP HC-EGFP HC-EGFP HC-EGFP HC-EGFP
103 2 HC-EGFP 200 HC-EGFP HC-EGFP HC-EGFP 200 HC-EGFP 200 HC-EGFP HC-EGFP HC-EGFP 200 g t P
104 %5 ml g 45 ml l 9 cm l 9 cm PTYG HC-EGFP HC-EGFP 200 HC-EGFP HC-EGFP
105 1 1 g x ± ± 0.36 HC-EGFP 2.46 ± ± 0.34 HC-EGFP ± ± 0.76 HC-EGFP HC-EGFP ± ± 0.49 HC-EGFP 1.86 ± ± 0.24 HC-EGFP ± ± 0.55 HC-EGFP HC-EGFP ± 0.79 t P g x ± ± 0.19 HC-EGFP 1.50 ± ± 0.45 HC-EGFP ± ± 0.41 HC-EGFP HC-EGFP ± ± 0.53 HC-EGFP 1.31 ± ± 1. HC-EGFP ± ± 0.03 HC-EGFP HC-EGFP ± 0.21 t P
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