JP5745849B2 - アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 - Google Patents
アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 Download PDFInfo
- Publication number
- JP5745849B2 JP5745849B2 JP2010528175A JP2010528175A JP5745849B2 JP 5745849 B2 JP5745849 B2 JP 5745849B2 JP 2010528175 A JP2010528175 A JP 2010528175A JP 2010528175 A JP2010528175 A JP 2010528175A JP 5745849 B2 JP5745849 B2 JP 5745849B2
- Authority
- JP
- Japan
- Prior art keywords
- ahas
- seq
- gene
- plant
- herbicide
- Prior art date
- Legal status (The legal status is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the status listed.)
- Active
Links
- 241000219198 Brassica Species 0.000 title claims description 28
- 108010000700 Acetolactate synthase Proteins 0.000 title claims description 26
- 235000011331 Brassica Nutrition 0.000 title claims description 23
- 108090000623 proteins and genes Proteins 0.000 claims description 229
- 230000035772 mutation Effects 0.000 claims description 198
- 241000196324 Embryophyta Species 0.000 claims description 172
- 238000006467 substitution reaction Methods 0.000 claims description 78
- 239000004009 herbicide Substances 0.000 claims description 77
- 102000004169 proteins and genes Human genes 0.000 claims description 66
- 238000000034 method Methods 0.000 claims description 65
- 230000002363 herbicidal effect Effects 0.000 claims description 64
- 235000018102 proteins Nutrition 0.000 claims description 64
- 125000003275 alpha amino acid group Chemical group 0.000 claims description 55
- 240000002791 Brassica napus Species 0.000 claims description 42
- QIVBCDIJIAJPQS-UHFFFAOYSA-N Tryptophan Natural products C1=CC=C2C(CC(N)C(O)=O)=CNC2=C1 QIVBCDIJIAJPQS-UHFFFAOYSA-N 0.000 claims description 35
- 230000008439 repair process Effects 0.000 claims description 30
- 230000002401 inhibitory effect Effects 0.000 claims description 27
- MTCFGRXMJLQNBG-UHFFFAOYSA-N Serine Natural products OCC(N)C(O)=O MTCFGRXMJLQNBG-UHFFFAOYSA-N 0.000 claims description 24
- 235000004977 Brassica sinapistrum Nutrition 0.000 claims description 21
- 125000000430 tryptophan group Chemical class [H]N([H])C(C(=O)O*)C([H])([H])C1=C([H])N([H])C2=C([H])C([H])=C([H])C([H])=C12 0.000 claims description 18
- QIVBCDIJIAJPQS-VIFPVBQESA-N L-tryptophane Chemical class C1=CC=C2C(C[C@H](N)C(O)=O)=CNC2=C1 QIVBCDIJIAJPQS-VIFPVBQESA-N 0.000 claims description 17
- 239000000203 mixture Substances 0.000 claims description 15
- ROHFNLRQFUQHCH-UHFFFAOYSA-N Leucine Natural products CC(C)CC(N)C(O)=O ROHFNLRQFUQHCH-UHFFFAOYSA-N 0.000 claims description 13
- 229940100389 Sulfonylurea Drugs 0.000 claims description 13
- CAAMSDWKXXPUJR-UHFFFAOYSA-N 3,5-dihydro-4H-imidazol-4-one Chemical class O=C1CNC=N1 CAAMSDWKXXPUJR-UHFFFAOYSA-N 0.000 claims description 12
- 230000009261 transgenic effect Effects 0.000 claims description 12
- 235000014698 Brassica juncea var multisecta Nutrition 0.000 claims description 11
- 235000006008 Brassica napus var napus Nutrition 0.000 claims description 11
- 235000006618 Brassica rapa subsp oleifera Nutrition 0.000 claims description 11
- ROHFNLRQFUQHCH-YFKPBYRVSA-N L-leucine Chemical compound CC(C)C[C@H](N)C(O)=O ROHFNLRQFUQHCH-YFKPBYRVSA-N 0.000 claims description 7
- YROXIXLRRCOBKF-UHFFFAOYSA-N sulfonylurea Chemical class OC(=N)N=S(=O)=O YROXIXLRRCOBKF-UHFFFAOYSA-N 0.000 claims description 7
- CDUVSERIDNVFDD-UHFFFAOYSA-N 2-pyrimidin-2-ylbenzenecarbothioic s-acid Chemical class OC(=S)C1=CC=CC=C1C1=NC=CC=N1 CDUVSERIDNVFDD-UHFFFAOYSA-N 0.000 claims description 6
- PRZRAMLXTKZUHF-UHFFFAOYSA-N 5-oxo-n-sulfonyl-4h-triazole-1-carboxamide Chemical class O=S(=O)=NC(=O)N1N=NCC1=O PRZRAMLXTKZUHF-UHFFFAOYSA-N 0.000 claims description 6
- 101150017040 I gene Proteins 0.000 claims description 6
- 230000003197 catalytic effect Effects 0.000 claims description 6
- 235000018417 cysteine Nutrition 0.000 claims description 6
- XUJNEKJLAYXESH-UHFFFAOYSA-N cysteine Natural products SCC(N)C(O)=O XUJNEKJLAYXESH-UHFFFAOYSA-N 0.000 claims description 6
- 125000001909 leucine group Chemical group [H]N(*)C(C(*)=O)C([H])([H])C(C([H])([H])[H])C([H])([H])[H] 0.000 claims description 6
- 229930182817 methionine Natural products 0.000 claims description 6
- 240000008042 Zea mays Species 0.000 claims description 5
- 241000219310 Beta vulgaris subsp. vulgaris Species 0.000 claims description 4
- FFEARJCKVFRZRR-BYPYZUCNSA-N L-methionine Chemical compound CSCC[C@H](N)C(O)=O FFEARJCKVFRZRR-BYPYZUCNSA-N 0.000 claims description 4
- 235000002637 Nicotiana tabacum Nutrition 0.000 claims description 4
- 244000061176 Nicotiana tabacum Species 0.000 claims description 4
- 235000021536 Sugar beet Nutrition 0.000 claims description 4
- 235000002017 Zea mays subsp mays Nutrition 0.000 claims description 4
- 230000005764 inhibitory process Effects 0.000 claims description 4
- 238000004519 manufacturing process Methods 0.000 claims description 4
- 230000008685 targeting Effects 0.000 claims description 4
- 230000008569 process Effects 0.000 claims description 3
- 244000291564 Allium cepa Species 0.000 claims description 2
- 235000002732 Allium cepa var. cepa Nutrition 0.000 claims description 2
- 244000144730 Amygdalus persica Species 0.000 claims description 2
- 244000075850 Avena orientalis Species 0.000 claims description 2
- 235000007319 Avena orientalis Nutrition 0.000 claims description 2
- 240000007124 Brassica oleracea Species 0.000 claims description 2
- 235000003899 Brassica oleracea var acephala Nutrition 0.000 claims description 2
- 235000011301 Brassica oleracea var capitata Nutrition 0.000 claims description 2
- 235000001169 Brassica oleracea var oleracea Nutrition 0.000 claims description 2
- 235000004936 Bromus mango Nutrition 0.000 claims description 2
- 235000005881 Calendula officinalis Nutrition 0.000 claims description 2
- 244000025254 Cannabis sativa Species 0.000 claims description 2
- 240000004160 Capsicum annuum Species 0.000 claims description 2
- 235000008534 Capsicum annuum var annuum Nutrition 0.000 claims description 2
- 235000005633 Chrysanthemum balsamita Nutrition 0.000 claims description 2
- 244000260524 Chrysanthemum balsamita Species 0.000 claims description 2
- 241000207199 Citrus Species 0.000 claims description 2
- 229920000742 Cotton Polymers 0.000 claims description 2
- 241000219112 Cucumis Species 0.000 claims description 2
- 235000015510 Cucumis melo subsp melo Nutrition 0.000 claims description 2
- 240000008067 Cucumis sativus Species 0.000 claims description 2
- 235000010799 Cucumis sativus var sativus Nutrition 0.000 claims description 2
- 235000002767 Daucus carota Nutrition 0.000 claims description 2
- 244000000626 Daucus carota Species 0.000 claims description 2
- 235000009355 Dianthus caryophyllus Nutrition 0.000 claims description 2
- 240000006497 Dianthus caryophyllus Species 0.000 claims description 2
- 235000016623 Fragaria vesca Nutrition 0.000 claims description 2
- 240000009088 Fragaria x ananassa Species 0.000 claims description 2
- 235000011363 Fragaria x ananassa Nutrition 0.000 claims description 2
- 244000068988 Glycine max Species 0.000 claims description 2
- 241000219146 Gossypium Species 0.000 claims description 2
- 244000020551 Helianthus annuus Species 0.000 claims description 2
- 235000003222 Helianthus annuus Nutrition 0.000 claims description 2
- 240000005979 Hordeum vulgare Species 0.000 claims description 2
- 235000007340 Hordeum vulgare Nutrition 0.000 claims description 2
- 240000001549 Ipomoea eriocarpa Species 0.000 claims description 2
- 235000005146 Ipomoea eriocarpa Nutrition 0.000 claims description 2
- 235000003228 Lactuca sativa Nutrition 0.000 claims description 2
- 240000008415 Lactuca sativa Species 0.000 claims description 2
- 241000234435 Lilium Species 0.000 claims description 2
- 235000004431 Linum usitatissimum Nutrition 0.000 claims description 2
- 240000006240 Linum usitatissimum Species 0.000 claims description 2
- 235000007688 Lycopersicon esculentum Nutrition 0.000 claims description 2
- 241000220225 Malus Species 0.000 claims description 2
- 235000011430 Malus pumila Nutrition 0.000 claims description 2
- 235000015103 Malus silvestris Nutrition 0.000 claims description 2
- 235000014826 Mangifera indica Nutrition 0.000 claims description 2
- 240000007228 Mangifera indica Species 0.000 claims description 2
- 240000004658 Medicago sativa Species 0.000 claims description 2
- 235000017587 Medicago sativa ssp. sativa Nutrition 0.000 claims description 2
- 240000005561 Musa balbisiana Species 0.000 claims description 2
- 235000018290 Musa x paradisiaca Nutrition 0.000 claims description 2
- 240000002853 Nelumbo nucifera Species 0.000 claims description 2
- 235000006508 Nelumbo nucifera Nutrition 0.000 claims description 2
- 235000006510 Nelumbo pentapetala Nutrition 0.000 claims description 2
- 240000007594 Oryza sativa Species 0.000 claims description 2
- 235000007164 Oryza sativa Nutrition 0.000 claims description 2
- 240000004713 Pisum sativum Species 0.000 claims description 2
- 235000010582 Pisum sativum Nutrition 0.000 claims description 2
- 241000219000 Populus Species 0.000 claims description 2
- 235000006040 Prunus persica var persica Nutrition 0.000 claims description 2
- 235000014443 Pyrus communis Nutrition 0.000 claims description 2
- 240000001987 Pyrus communis Species 0.000 claims description 2
- 240000000111 Saccharum officinarum Species 0.000 claims description 2
- 235000007201 Saccharum officinarum Nutrition 0.000 claims description 2
- 235000007238 Secale cereale Nutrition 0.000 claims description 2
- 240000003768 Solanum lycopersicum Species 0.000 claims description 2
- 235000002597 Solanum melongena Nutrition 0.000 claims description 2
- 244000061458 Solanum melongena Species 0.000 claims description 2
- 235000002595 Solanum tuberosum Nutrition 0.000 claims description 2
- 244000061456 Solanum tuberosum Species 0.000 claims description 2
- 240000003829 Sorghum propinquum Species 0.000 claims description 2
- 235000011684 Sorghum saccharatum Nutrition 0.000 claims description 2
- 235000009184 Spondias indica Nutrition 0.000 claims description 2
- 240000000785 Tagetes erecta Species 0.000 claims description 2
- 235000021307 Triticum Nutrition 0.000 claims description 2
- 244000098338 Triticum aestivum Species 0.000 claims description 2
- 241000722921 Tulipa gesneriana Species 0.000 claims description 2
- 235000010749 Vicia faba Nutrition 0.000 claims description 2
- 240000006677 Vicia faba Species 0.000 claims description 2
- 235000002098 Vicia faba var. major Nutrition 0.000 claims description 2
- 235000009754 Vitis X bourquina Nutrition 0.000 claims description 2
- 235000012333 Vitis X labruscana Nutrition 0.000 claims description 2
- 240000006365 Vitis vinifera Species 0.000 claims description 2
- 235000014787 Vitis vinifera Nutrition 0.000 claims description 2
- 235000005824 Zea mays ssp. parviglumis Nutrition 0.000 claims description 2
- FJJCIZWZNKZHII-UHFFFAOYSA-N [4,6-bis(cyanoamino)-1,3,5-triazin-2-yl]cyanamide Chemical compound N#CNC1=NC(NC#N)=NC(NC#N)=N1 FJJCIZWZNKZHII-UHFFFAOYSA-N 0.000 claims description 2
- 235000020971 citrus fruits Nutrition 0.000 claims description 2
- 235000005822 corn Nutrition 0.000 claims description 2
- 239000004459 forage Substances 0.000 claims description 2
- 235000009566 rice Nutrition 0.000 claims description 2
- 244000188595 Brassica sinapistrum Species 0.000 claims 1
- 235000009811 Momordica charantia Nutrition 0.000 claims 1
- 241000209056 Secale Species 0.000 claims 1
- 235000008322 Trichosanthes cucumerina Nutrition 0.000 claims 1
- 244000078912 Trichosanthes cucumerina Species 0.000 claims 1
- 210000000554 iris Anatomy 0.000 claims 1
- 239000002773 nucleotide Substances 0.000 description 152
- 125000003729 nucleotide group Chemical group 0.000 description 152
- 210000004027 cell Anatomy 0.000 description 55
- 235000001014 amino acid Nutrition 0.000 description 52
- 150000007523 nucleic acids Chemical class 0.000 description 39
- 102000039446 nucleic acids Human genes 0.000 description 38
- 108020004707 nucleic acids Proteins 0.000 description 38
- 229940024606 amino acid Drugs 0.000 description 36
- 150000001413 amino acids Chemical class 0.000 description 36
- 108091034117 Oligonucleotide Proteins 0.000 description 35
- 235000011293 Brassica napus Nutrition 0.000 description 32
- 210000001938 protoplast Anatomy 0.000 description 31
- 108091032973 (ribonucleotides)n+m Proteins 0.000 description 30
- 108020004414 DNA Proteins 0.000 description 30
- 210000001519 tissue Anatomy 0.000 description 25
- 206010020649 Hyperkeratosis Diseases 0.000 description 21
- 239000002609 medium Substances 0.000 description 20
- 230000000295 complement effect Effects 0.000 description 19
- 239000000047 product Substances 0.000 description 17
- 235000004279 alanine Nutrition 0.000 description 16
- QNAYBMKLOCPYGJ-REOHCLBHSA-N L-alanine Chemical compound C[C@H](N)C(O)=O QNAYBMKLOCPYGJ-REOHCLBHSA-N 0.000 description 14
- JLCPHMBAVCMARE-UHFFFAOYSA-N [3-[[3-[[3-[[3-[[3-[[3-[[3-[[3-[[3-[[3-[[3-[[5-(2-amino-6-oxo-1H-purin-9-yl)-3-[[3-[[3-[[3-[[3-[[3-[[5-(2-amino-6-oxo-1H-purin-9-yl)-3-[[5-(2-amino-6-oxo-1H-purin-9-yl)-3-hydroxyoxolan-2-yl]methoxy-hydroxyphosphoryl]oxyoxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(5-methyl-2,4-dioxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxyoxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(5-methyl-2,4-dioxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(4-amino-2-oxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(5-methyl-2,4-dioxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(5-methyl-2,4-dioxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(4-amino-2-oxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(4-amino-2-oxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(4-amino-2-oxopyrimidin-1-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(6-aminopurin-9-yl)oxolan-2-yl]methoxy-hydroxyphosphoryl]oxy-5-(4-amino-2-oxopyrimidin-1-yl)oxolan-2-yl]methyl [5-(6-aminopurin-9-yl)-2-(hydroxymethyl)oxolan-3-yl] hydrogen phosphate Polymers Cc1cn(C2CC(OP(O)(=O)OCC3OC(CC3OP(O)(=O)OCC3OC(CC3O)n3cnc4c3nc(N)[nH]c4=O)n3cnc4c3nc(N)[nH]c4=O)C(COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3COP(O)(=O)OC3CC(OC3CO)n3cnc4c(N)ncnc34)n3ccc(N)nc3=O)n3cnc4c(N)ncnc34)n3ccc(N)nc3=O)n3ccc(N)nc3=O)n3ccc(N)nc3=O)n3cnc4c(N)ncnc34)n3cnc4c(N)ncnc34)n3cc(C)c(=O)[nH]c3=O)n3cc(C)c(=O)[nH]c3=O)n3ccc(N)nc3=O)n3cc(C)c(=O)[nH]c3=O)n3cnc4c3nc(N)[nH]c4=O)n3cnc4c(N)ncnc34)n3cnc4c(N)ncnc34)n3cnc4c(N)ncnc34)n3cnc4c(N)ncnc34)O2)c(=O)[nH]c1=O JLCPHMBAVCMARE-UHFFFAOYSA-N 0.000 description 14
- 125000001424 substituent group Chemical group 0.000 description 13
- 235000003704 aspartic acid Nutrition 0.000 description 12
- CKLJMWTZIZZHCS-REOHCLBHSA-N aspartic acid group Chemical group N[C@@H](CC(=O)O)C(=O)O CKLJMWTZIZZHCS-REOHCLBHSA-N 0.000 description 12
- OQFSQFPPLPISGP-UHFFFAOYSA-N beta-carboxyaspartic acid Natural products OC(=O)C(N)C(C(O)=O)C(O)=O OQFSQFPPLPISGP-UHFFFAOYSA-N 0.000 description 12
- 239000012634 fragment Substances 0.000 description 12
- 240000000385 Brassica napus var. napus Species 0.000 description 11
- 229920001223 polyethylene glycol Polymers 0.000 description 11
- 230000014616 translation Effects 0.000 description 11
- DCXYFEDJOCDNAF-UHFFFAOYSA-N Asparagine Natural products OC(=O)C(N)CC(N)=O DCXYFEDJOCDNAF-UHFFFAOYSA-N 0.000 description 10
- 239000002202 Polyethylene glycol Substances 0.000 description 10
- 238000007844 allele-specific PCR Methods 0.000 description 10
- 235000009582 asparagine Nutrition 0.000 description 10
- 229960001230 asparagine Drugs 0.000 description 10
- 210000000056 organ Anatomy 0.000 description 10
- 238000012360 testing method Methods 0.000 description 10
- 108700028369 Alleles Proteins 0.000 description 9
- 108091028043 Nucleic acid sequence Proteins 0.000 description 9
- 108090000765 processed proteins & peptides Proteins 0.000 description 9
- 239000007921 spray Substances 0.000 description 9
- 238000013519 translation Methods 0.000 description 9
- AYFVYJQAPQTCCC-UHFFFAOYSA-N Threonine Natural products CC(O)C(N)C(O)=O AYFVYJQAPQTCCC-UHFFFAOYSA-N 0.000 description 8
- 239000004473 Threonine Substances 0.000 description 8
- KZSNJWFQEVHDMF-UHFFFAOYSA-N Valine Natural products CC(C)C(N)C(O)=O KZSNJWFQEVHDMF-UHFFFAOYSA-N 0.000 description 8
- 230000008859 change Effects 0.000 description 8
- 241000894007 species Species 0.000 description 8
- 239000004474 valine Substances 0.000 description 8
- FWMNVWWHGCHHJJ-SKKKGAJSSA-N 4-amino-1-[(2r)-6-amino-2-[[(2r)-2-[[(2r)-2-[[(2r)-2-amino-3-phenylpropanoyl]amino]-3-phenylpropanoyl]amino]-4-methylpentanoyl]amino]hexanoyl]piperidine-4-carboxylic acid Chemical compound C([C@H](C(=O)N[C@H](CC(C)C)C(=O)N[C@H](CCCCN)C(=O)N1CCC(N)(CC1)C(O)=O)NC(=O)[C@H](N)CC=1C=CC=CC=1)C1=CC=CC=C1 FWMNVWWHGCHHJJ-SKKKGAJSSA-N 0.000 description 7
- LFQSCWFLJHTTHZ-UHFFFAOYSA-N Ethanol Chemical compound CCO LFQSCWFLJHTTHZ-UHFFFAOYSA-N 0.000 description 7
- 238000011161 development Methods 0.000 description 7
- 230000018109 developmental process Effects 0.000 description 7
- 229920001184 polypeptide Polymers 0.000 description 7
- 102000004196 processed proteins & peptides Human genes 0.000 description 7
- 238000012163 sequencing technique Methods 0.000 description 7
- 125000002987 valine group Chemical group [H]N([H])C([H])(C(*)=O)C([H])(C([H])([H])[H])C([H])([H])[H] 0.000 description 7
- 108091026890 Coding region Proteins 0.000 description 6
- 229920001410 Microfiber Polymers 0.000 description 6
- 108091028664 Ribonucleotide Proteins 0.000 description 6
- 238000012300 Sequence Analysis Methods 0.000 description 6
- 125000000613 asparagine group Chemical group N[C@@H](CC(N)=O)C(=O)* 0.000 description 6
- 239000002299 complementary DNA Substances 0.000 description 6
- 239000000975 dye Substances 0.000 description 6
- 125000005647 linker group Chemical group 0.000 description 6
- 239000003658 microfiber Substances 0.000 description 6
- 238000002703 mutagenesis Methods 0.000 description 6
- 231100000350 mutagenesis Toxicity 0.000 description 6
- 229940046166 oligodeoxynucleotide Drugs 0.000 description 6
- 239000013612 plasmid Substances 0.000 description 6
- 239000002336 ribonucleotide Substances 0.000 description 6
- 125000002652 ribonucleotide group Chemical group 0.000 description 6
- ATHGHQPFGPMSJY-UHFFFAOYSA-N spermidine Chemical compound NCCCCNCCCN ATHGHQPFGPMSJY-UHFFFAOYSA-N 0.000 description 6
- 125000000341 threoninyl group Chemical group [H]OC([H])(C([H])([H])[H])C([H])(N([H])[H])C(*)=O 0.000 description 6
- RWQNBRDOKXIBIV-UHFFFAOYSA-N thymine Chemical compound CC1=CNC(=O)NC1=O RWQNBRDOKXIBIV-UHFFFAOYSA-N 0.000 description 6
- 238000011282 treatment Methods 0.000 description 6
- 241000219194 Arabidopsis Species 0.000 description 5
- 108020004705 Codon Proteins 0.000 description 5
- 230000000903 blocking effect Effects 0.000 description 5
- 238000003780 insertion Methods 0.000 description 5
- 230000037431 insertion Effects 0.000 description 5
- 238000012986 modification Methods 0.000 description 5
- 230000004048 modification Effects 0.000 description 5
- 239000000243 solution Substances 0.000 description 5
- 239000000126 substance Substances 0.000 description 5
- NUPJIGQFXCQJBK-UHFFFAOYSA-N 2-(4-isopropyl-4-methyl-5-oxo-4,5-dihydro-1H-imidazol-2-yl)-5-(methoxymethyl)nicotinic acid Chemical compound OC(=O)C1=CC(COC)=CN=C1C1=NC(C)(C(C)C)C(=O)N1 NUPJIGQFXCQJBK-UHFFFAOYSA-N 0.000 description 4
- KDCGOANMDULRCW-UHFFFAOYSA-N 7H-purine Chemical compound N1=CNC2=NC=NC2=C1 KDCGOANMDULRCW-UHFFFAOYSA-N 0.000 description 4
- 101100179609 Arabidopsis thaliana ALS gene Proteins 0.000 description 4
- 239000004475 Arginine Substances 0.000 description 4
- 102000004190 Enzymes Human genes 0.000 description 4
- 108090000790 Enzymes Proteins 0.000 description 4
- WHUUTDBJXJRKMK-UHFFFAOYSA-N Glutamic acid Natural products OC(=O)C(N)CCC(O)=O WHUUTDBJXJRKMK-UHFFFAOYSA-N 0.000 description 4
- 101150098499 III gene Proteins 0.000 description 4
- 239000005566 Imazamox Substances 0.000 description 4
- DCXYFEDJOCDNAF-REOHCLBHSA-N L-asparagine Chemical compound OC(=O)[C@@H](N)CC(N)=O DCXYFEDJOCDNAF-REOHCLBHSA-N 0.000 description 4
- 239000005586 Nicosulfuron Substances 0.000 description 4
- ODKSFYDXXFIFQN-UHFFFAOYSA-N arginine Natural products OC(=O)C(N)CCCNC(N)=N ODKSFYDXXFIFQN-UHFFFAOYSA-N 0.000 description 4
- OPTASPLRGRRNAP-UHFFFAOYSA-N cytosine Chemical compound NC=1C=CNC(=O)N=1 OPTASPLRGRRNAP-UHFFFAOYSA-N 0.000 description 4
- 238000012217 deletion Methods 0.000 description 4
- 230000037430 deletion Effects 0.000 description 4
- 238000004520 electroporation Methods 0.000 description 4
- 229940088598 enzyme Drugs 0.000 description 4
- 230000014509 gene expression Effects 0.000 description 4
- 235000013922 glutamic acid Nutrition 0.000 description 4
- 239000004220 glutamic acid Substances 0.000 description 4
- UYTPUPDQBNUYGX-UHFFFAOYSA-N guanine Chemical compound O=C1NC(N)=NC2=C1N=CN2 UYTPUPDQBNUYGX-UHFFFAOYSA-N 0.000 description 4
- 239000000463 material Substances 0.000 description 4
- 230000001404 mediated effect Effects 0.000 description 4
- 238000000520 microinjection Methods 0.000 description 4
- RTCOGUMHFFWOJV-UHFFFAOYSA-N nicosulfuron Chemical compound COC1=CC(OC)=NC(NC(=O)NS(=O)(=O)C=2C(=CC=CN=2)C(=O)N(C)C)=N1 RTCOGUMHFFWOJV-UHFFFAOYSA-N 0.000 description 4
- 239000002777 nucleoside Substances 0.000 description 4
- 238000000746 purification Methods 0.000 description 4
- 239000006152 selective media Substances 0.000 description 4
- 239000000725 suspension Substances 0.000 description 4
- LOQQVLXUKHKNIA-UHFFFAOYSA-N thifensulfuron Chemical compound COC1=NC(C)=NC(NC(=O)NS(=O)(=O)C2=C(SC=C2)C(O)=O)=N1 LOQQVLXUKHKNIA-UHFFFAOYSA-N 0.000 description 4
- 230000009466 transformation Effects 0.000 description 4
- 239000013598 vector Substances 0.000 description 4
- XLYOFNOQVPJJNP-UHFFFAOYSA-N water Chemical compound O XLYOFNOQVPJJNP-UHFFFAOYSA-N 0.000 description 4
- 229930024421 Adenine Natural products 0.000 description 3
- GFFGJBXGBJISGV-UHFFFAOYSA-N Adenine Chemical compound NC1=NC=NC2=C1N=CN2 GFFGJBXGBJISGV-UHFFFAOYSA-N 0.000 description 3
- PEDCQBHIVMGVHV-UHFFFAOYSA-N Glycerine Chemical compound OCC(O)CO PEDCQBHIVMGVHV-UHFFFAOYSA-N 0.000 description 3
- 108091005461 Nucleic proteins Proteins 0.000 description 3
- OAICVXFJPJFONN-UHFFFAOYSA-N Phosphorus Chemical compound [P] OAICVXFJPJFONN-UHFFFAOYSA-N 0.000 description 3
- CZPWVGJYEJSRLH-UHFFFAOYSA-N Pyrimidine Chemical compound C1=CN=CN=C1 CZPWVGJYEJSRLH-UHFFFAOYSA-N 0.000 description 3
- 239000005626 Tribenuron Substances 0.000 description 3
- 229960000643 adenine Drugs 0.000 description 3
- 235000010443 alginic acid Nutrition 0.000 description 3
- 229920000615 alginic acid Polymers 0.000 description 3
- 210000002421 cell wall Anatomy 0.000 description 3
- 230000001413 cellular effect Effects 0.000 description 3
- 230000006870 function Effects 0.000 description 3
- 238000000338 in vitro Methods 0.000 description 3
- 239000004137 magnesium phosphate Substances 0.000 description 3
- 108020004999 messenger RNA Proteins 0.000 description 3
- 125000003835 nucleoside group Chemical group 0.000 description 3
- 150000004713 phosphodiesters Chemical class 0.000 description 3
- 150000008300 phosphoramidites Chemical class 0.000 description 3
- 229910052698 phosphorus Inorganic materials 0.000 description 3
- 239000011574 phosphorus Substances 0.000 description 3
- 229920000642 polymer Polymers 0.000 description 3
- 229940063673 spermidine Drugs 0.000 description 3
- 229940113082 thymine Drugs 0.000 description 3
- BQZXUHDXIARLEO-UHFFFAOYSA-N tribenuron Chemical compound COC1=NC(C)=NC(N(C)C(=O)NS(=O)(=O)C=2C(=CC=CC=2)C(O)=O)=N1 BQZXUHDXIARLEO-UHFFFAOYSA-N 0.000 description 3
- YBJHBAHKTGYVGT-ZKWXMUAHSA-N (+)-Biotin Chemical compound N1C(=O)N[C@@H]2[C@H](CCCCC(=O)O)SC[C@@H]21 YBJHBAHKTGYVGT-ZKWXMUAHSA-N 0.000 description 2
- 108091093088 Amplicon Proteins 0.000 description 2
- OKTJSMMVPCPJKN-UHFFFAOYSA-N Carbon Chemical compound [C] OKTJSMMVPCPJKN-UHFFFAOYSA-N 0.000 description 2
- 108091033380 Coding strand Proteins 0.000 description 2
- 239000005562 Glyphosate Substances 0.000 description 2
- YNAVUWVOSKDBBP-UHFFFAOYSA-N Morpholine Chemical compound C1COCCN1 YNAVUWVOSKDBBP-UHFFFAOYSA-N 0.000 description 2
- 102000006382 Ribonucleases Human genes 0.000 description 2
- 108010083644 Ribonucleases Proteins 0.000 description 2
- 108020004566 Transfer RNA Proteins 0.000 description 2
- ISAKRJDGNUQOIC-UHFFFAOYSA-N Uracil Chemical compound O=C1C=CNC(=O)N1 ISAKRJDGNUQOIC-UHFFFAOYSA-N 0.000 description 2
- 235000016383 Zea mays subsp huehuetenangensis Nutrition 0.000 description 2
- 239000004480 active ingredient Substances 0.000 description 2
- 150000001294 alanine derivatives Chemical class 0.000 description 2
- 239000000783 alginic acid Substances 0.000 description 2
- 229960001126 alginic acid Drugs 0.000 description 2
- 150000004781 alginic acids Chemical class 0.000 description 2
- 230000003321 amplification Effects 0.000 description 2
- 230000000692 anti-sense effect Effects 0.000 description 2
- 238000013459 approach Methods 0.000 description 2
- 239000000648 calcium alginate Substances 0.000 description 2
- 235000010410 calcium alginate Nutrition 0.000 description 2
- 229960002681 calcium alginate Drugs 0.000 description 2
- OKHHGHGGPDJQHR-YMOPUZKJSA-L calcium;(2s,3s,4s,5s,6r)-6-[(2r,3s,4r,5s,6r)-2-carboxy-6-[(2r,3s,4r,5s,6r)-2-carboxylato-4,5,6-trihydroxyoxan-3-yl]oxy-4,5-dihydroxyoxan-3-yl]oxy-3,4,5-trihydroxyoxane-2-carboxylate Chemical compound [Ca+2].O[C@@H]1[C@H](O)[C@H](O)O[C@@H](C([O-])=O)[C@H]1O[C@H]1[C@@H](O)[C@@H](O)[C@H](O[C@H]2[C@H]([C@@H](O)[C@H](O)[C@H](O2)C([O-])=O)O)[C@H](C(O)=O)O1 OKHHGHGGPDJQHR-YMOPUZKJSA-L 0.000 description 2
- 229910052799 carbon Inorganic materials 0.000 description 2
- 230000032823 cell division Effects 0.000 description 2
- 238000006243 chemical reaction Methods 0.000 description 2
- 239000003153 chemical reaction reagent Substances 0.000 description 2
- 125000001309 chloro group Chemical group Cl* 0.000 description 2
- HVYWMOMLDIMFJA-DPAQBDIFSA-N cholesterol Chemical compound C1C=C2C[C@@H](O)CC[C@]2(C)[C@@H]2[C@@H]1[C@@H]1CC[C@H]([C@H](C)CCCC(C)C)[C@@]1(C)CC2 HVYWMOMLDIMFJA-DPAQBDIFSA-N 0.000 description 2
- 230000002759 chromosomal effect Effects 0.000 description 2
- 150000001875 compounds Chemical class 0.000 description 2
- 230000001276 controlling effect Effects 0.000 description 2
- 229940104302 cytosine Drugs 0.000 description 2
- 231100000673 dose–response relationship Toxicity 0.000 description 2
- 235000013399 edible fruits Nutrition 0.000 description 2
- 230000000694 effects Effects 0.000 description 2
- 210000002257 embryonic structure Anatomy 0.000 description 2
- 230000002255 enzymatic effect Effects 0.000 description 2
- 230000007717 exclusion Effects 0.000 description 2
- 238000002474 experimental method Methods 0.000 description 2
- 239000013604 expression vector Substances 0.000 description 2
- 239000000835 fiber Substances 0.000 description 2
- 238000010363 gene targeting Methods 0.000 description 2
- 230000002068 genetic effect Effects 0.000 description 2
- 235000003869 genetically modified organism Nutrition 0.000 description 2
- XDDAORKBJWWYJS-UHFFFAOYSA-N glyphosate Chemical compound OC(=O)CNCP(O)(O)=O XDDAORKBJWWYJS-UHFFFAOYSA-N 0.000 description 2
- 229940097068 glyphosate Drugs 0.000 description 2
- 239000001963 growth medium Substances 0.000 description 2
- 238000009396 hybridization Methods 0.000 description 2
- 230000006872 improvement Effects 0.000 description 2
- 238000009830 intercalation Methods 0.000 description 2
- 238000002955 isolation Methods 0.000 description 2
- 230000000670 limiting effect Effects 0.000 description 2
- 239000007788 liquid Substances 0.000 description 2
- 235000009973 maize Nutrition 0.000 description 2
- 230000013011 mating Effects 0.000 description 2
- 125000001360 methionine group Chemical group N[C@@H](CCSC)C(=O)* 0.000 description 2
- 229910052757 nitrogen Inorganic materials 0.000 description 2
- 238000003199 nucleic acid amplification method Methods 0.000 description 2
- 238000002360 preparation method Methods 0.000 description 2
- 238000001243 protein synthesis Methods 0.000 description 2
- 150000003230 pyrimidines Chemical class 0.000 description 2
- 230000001105 regulatory effect Effects 0.000 description 2
- 238000011160 research Methods 0.000 description 2
- 238000012216 screening Methods 0.000 description 2
- 238000012546 transfer Methods 0.000 description 2
- PRPINYUDVPFIRX-UHFFFAOYSA-N 1-naphthaleneacetic acid Chemical compound C1=CC=C2C(CC(=O)O)=CC=CC2=C1 PRPINYUDVPFIRX-UHFFFAOYSA-N 0.000 description 1
- SXGZJKUKBWWHRA-UHFFFAOYSA-N 2-(N-morpholiniumyl)ethanesulfonate Chemical compound [O-]S(=O)(=O)CC[NH+]1CCOCC1 SXGZJKUKBWWHRA-UHFFFAOYSA-N 0.000 description 1
- 108010020183 3-phosphoshikimate 1-carboxyvinyltransferase Proteins 0.000 description 1
- ZXICIFDEGMIMJY-UHFFFAOYSA-N 4-[2-[3-[3,3-dimethyl-1-(4-sulfobutyl)indol-2-ylidene]prop-1-enyl]-3,3-dimethylindol-1-ium-1-yl]butane-1-sulfonate Chemical class OS(=O)(=O)CCCCN1C2=CC=CC=C2C(C)(C)\C1=C\C=C\C1=[N+](CCCCS([O-])(=O)=O)C2=CC=CC=C2C1(C)C ZXICIFDEGMIMJY-UHFFFAOYSA-N 0.000 description 1
- FHVDTGUDJYJELY-UHFFFAOYSA-N 6-{[2-carboxy-4,5-dihydroxy-6-(phosphanyloxy)oxan-3-yl]oxy}-4,5-dihydroxy-3-phosphanyloxane-2-carboxylic acid Chemical compound O1C(C(O)=O)C(P)C(O)C(O)C1OC1C(C(O)=O)OC(OP)C(O)C1O FHVDTGUDJYJELY-UHFFFAOYSA-N 0.000 description 1
- PVFOHMXILQEIHX-UHFFFAOYSA-N 8-[(6-bromo-1,3-benzodioxol-5-yl)sulfanyl]-9-[2-(2-bromophenyl)ethyl]purin-6-amine Chemical compound C=1C=2OCOC=2C=C(Br)C=1SC1=NC=2C(N)=NC=NC=2N1CCC1=CC=CC=C1Br PVFOHMXILQEIHX-UHFFFAOYSA-N 0.000 description 1
- 235000007173 Abies balsamea Nutrition 0.000 description 1
- 229920000936 Agarose Polymers 0.000 description 1
- 108090000668 Annexin A2 Proteins 0.000 description 1
- 102100034613 Annexin A2 Human genes 0.000 description 1
- 108090000656 Annexin A6 Proteins 0.000 description 1
- 102100034278 Annexin A6 Human genes 0.000 description 1
- 239000004857 Balsam Substances 0.000 description 1
- 241000335053 Beta vulgaris Species 0.000 description 1
- 108091003079 Bovine Serum Albumin Proteins 0.000 description 1
- 241000219193 Brassicaceae Species 0.000 description 1
- 108010059892 Cellulase Proteins 0.000 description 1
- 241000218631 Coniferophyta Species 0.000 description 1
- 244000301850 Cupressus sempervirens Species 0.000 description 1
- FBPFZTCFMRRESA-KVTDHHQDSA-N D-Mannitol Chemical compound OC[C@@H](O)[C@@H](O)[C@H](O)[C@H](O)CO FBPFZTCFMRRESA-KVTDHHQDSA-N 0.000 description 1
- 238000010442 DNA editing Methods 0.000 description 1
- 101150111720 EPSPS gene Proteins 0.000 description 1
- 108700024394 Exon Proteins 0.000 description 1
- 108091027305 Heteroduplex Proteins 0.000 description 1
- 101150062179 II gene Proteins 0.000 description 1
- 244000018716 Impatiens biflora Species 0.000 description 1
- 108091092195 Intron Proteins 0.000 description 1
- KZSNJWFQEVHDMF-BYPYZUCNSA-N L-valine Chemical compound CC(C)[C@H](N)C(O)=O KZSNJWFQEVHDMF-BYPYZUCNSA-N 0.000 description 1
- 229930195725 Mannitol Natural products 0.000 description 1
- 108091027974 Mature messenger RNA Proteins 0.000 description 1
- 108010021466 Mutant Proteins Proteins 0.000 description 1
- 102000008300 Mutant Proteins Human genes 0.000 description 1
- 206010028980 Neoplasm Diseases 0.000 description 1
- 241000208125 Nicotiana Species 0.000 description 1
- 240000002582 Oryza sativa Indica Group Species 0.000 description 1
- 238000010222 PCR analysis Methods 0.000 description 1
- 229910019142 PO4 Inorganic materials 0.000 description 1
- 108091093037 Peptide nucleic acid Proteins 0.000 description 1
- GPGLBXMQFQQXDV-UHFFFAOYSA-N Primisulfuron Chemical compound OC(=O)C1=CC=CC=C1S(=O)(=O)NC(=O)NC1=NC(OC(F)F)=CC(OC(F)F)=N1 GPGLBXMQFQQXDV-UHFFFAOYSA-N 0.000 description 1
- 108010039491 Ricin Proteins 0.000 description 1
- 239000005616 Rimsulfuron Substances 0.000 description 1
- 244000082988 Secale cereale Species 0.000 description 1
- 244000204900 Talipariti tiliaceum Species 0.000 description 1
- 241000607479 Yersinia pestis Species 0.000 description 1
- 235000007244 Zea mays Nutrition 0.000 description 1
- 108010017070 Zinc Finger Nucleases Proteins 0.000 description 1
- WNDWKKPBLAKXMI-UHFFFAOYSA-N [5-(benzenesulfonyl)-2-nitrophenyl] 4-chlorobenzoate Chemical compound [O-][N+](=O)C1=CC=C(S(=O)(=O)C=2C=CC=CC=2)C=C1OC(=O)C1=CC=C(Cl)C=C1 WNDWKKPBLAKXMI-UHFFFAOYSA-N 0.000 description 1
- 239000002253 acid Substances 0.000 description 1
- 239000003905 agrochemical Substances 0.000 description 1
- 229940072056 alginate Drugs 0.000 description 1
- 125000003545 alkoxy group Chemical group 0.000 description 1
- 230000004075 alteration Effects 0.000 description 1
- 125000000539 amino acid group Chemical group 0.000 description 1
- 238000000137 annealing Methods 0.000 description 1
- 125000004429 atom Chemical group 0.000 description 1
- 230000001580 bacterial effect Effects 0.000 description 1
- 230000037429 base substitution Effects 0.000 description 1
- 230000008901 benefit Effects 0.000 description 1
- 235000020958 biotin Nutrition 0.000 description 1
- 229960002685 biotin Drugs 0.000 description 1
- 239000011616 biotin Substances 0.000 description 1
- 229940098773 bovine serum albumin Drugs 0.000 description 1
- 238000009395 breeding Methods 0.000 description 1
- 230000001488 breeding effect Effects 0.000 description 1
- 125000001246 bromo group Chemical group Br* 0.000 description 1
- 210000004899 c-terminal region Anatomy 0.000 description 1
- 125000002091 cationic group Chemical group 0.000 description 1
- 238000004113 cell culture Methods 0.000 description 1
- 210000000170 cell membrane Anatomy 0.000 description 1
- 230000010307 cell transformation Effects 0.000 description 1
- 229940106157 cellulase Drugs 0.000 description 1
- 239000001913 cellulose Substances 0.000 description 1
- 229920002678 cellulose Polymers 0.000 description 1
- 238000012512 characterization method Methods 0.000 description 1
- 235000012000 cholesterol Nutrition 0.000 description 1
- 238000010367 cloning Methods 0.000 description 1
- 230000002860 competitive effect Effects 0.000 description 1
- 238000009833 condensation Methods 0.000 description 1
- 230000005494 condensation Effects 0.000 description 1
- 235000009508 confectionery Nutrition 0.000 description 1
- 238000012790 confirmation Methods 0.000 description 1
- 230000021615 conjugation Effects 0.000 description 1
- 238000012937 correction Methods 0.000 description 1
- 210000000805 cytoplasm Anatomy 0.000 description 1
- 230000003111 delayed effect Effects 0.000 description 1
- 238000000432 density-gradient centrifugation Methods 0.000 description 1
- 239000005547 deoxyribonucleotide Substances 0.000 description 1
- 125000002637 deoxyribonucleotide group Chemical group 0.000 description 1
- 235000021186 dishes Nutrition 0.000 description 1
- 238000005516 engineering process Methods 0.000 description 1
- 239000003623 enhancer Substances 0.000 description 1
- 230000001747 exhibiting effect Effects 0.000 description 1
- 238000000605 extraction Methods 0.000 description 1
- 238000010304 firing Methods 0.000 description 1
- 239000007850 fluorescent dye Substances 0.000 description 1
- 125000001153 fluoro group Chemical group F* 0.000 description 1
- 230000004927 fusion Effects 0.000 description 1
- 238000012239 gene modification Methods 0.000 description 1
- 230000004077 genetic alteration Effects 0.000 description 1
- 231100000118 genetic alteration Toxicity 0.000 description 1
- 238000012252 genetic analysis Methods 0.000 description 1
- 230000035784 germination Effects 0.000 description 1
- 229910052736 halogen Chemical group 0.000 description 1
- 150000002367 halogens Chemical group 0.000 description 1
- 238000002744 homologous recombination Methods 0.000 description 1
- 230000006801 homologous recombination Effects 0.000 description 1
- 125000002887 hydroxy group Chemical group [H]O* 0.000 description 1
- 238000001727 in vivo Methods 0.000 description 1
- 238000011065 in-situ storage Methods 0.000 description 1
- 238000011534 incubation Methods 0.000 description 1
- 125000001041 indolyl group Chemical group 0.000 description 1
- 239000003112 inhibitor Substances 0.000 description 1
- 238000002347 injection Methods 0.000 description 1
- 239000007924 injection Substances 0.000 description 1
- 230000010354 integration Effects 0.000 description 1
- NBQNWMBBSKPBAY-UHFFFAOYSA-N iodixanol Chemical compound IC=1C(C(=O)NCC(O)CO)=C(I)C(C(=O)NCC(O)CO)=C(I)C=1N(C(=O)C)CC(O)CN(C(C)=O)C1=C(I)C(C(=O)NCC(O)CO)=C(I)C(C(=O)NCC(O)CO)=C1I NBQNWMBBSKPBAY-UHFFFAOYSA-N 0.000 description 1
- 229960004359 iodixanol Drugs 0.000 description 1
- 125000001449 isopropyl group Chemical group [H]C([H])([H])C([H])(*)C([H])([H])[H] 0.000 description 1
- 230000002147 killing effect Effects 0.000 description 1
- 230000003902 lesion Effects 0.000 description 1
- 239000000594 mannitol Substances 0.000 description 1
- 235000010355 mannitol Nutrition 0.000 description 1
- 230000007246 mechanism Effects 0.000 description 1
- 239000012528 membrane Substances 0.000 description 1
- 229910052751 metal Inorganic materials 0.000 description 1
- 239000002184 metal Substances 0.000 description 1
- MYWUZJCMWCOHBA-VIFPVBQESA-N methamphetamine Chemical compound CN[C@@H](C)CC1=CC=CC=C1 MYWUZJCMWCOHBA-VIFPVBQESA-N 0.000 description 1
- 125000000325 methylidene group Chemical group [H]C([H])=* 0.000 description 1
- 239000004005 microsphere Substances 0.000 description 1
- 238000002156 mixing Methods 0.000 description 1
- 239000003607 modifier Substances 0.000 description 1
- 238000007479 molecular analysis Methods 0.000 description 1
- 231100000219 mutagenic Toxicity 0.000 description 1
- 230000003505 mutagenic effect Effects 0.000 description 1
- 230000001338 necrotic effect Effects 0.000 description 1
- 231100000252 nontoxic Toxicity 0.000 description 1
- 230000003000 nontoxic effect Effects 0.000 description 1
- 238000009377 nuclear transmutation Methods 0.000 description 1
- 150000003833 nucleoside derivatives Chemical class 0.000 description 1
- 210000001672 ovary Anatomy 0.000 description 1
- 230000036961 partial effect Effects 0.000 description 1
- 230000035515 penetration Effects 0.000 description 1
- 239000010452 phosphate Substances 0.000 description 1
- 125000002467 phosphate group Chemical group [H]OP(=O)(O[H])O[*] 0.000 description 1
- 210000000745 plant chromosome Anatomy 0.000 description 1
- 210000002706 plastid Anatomy 0.000 description 1
- 238000003752 polymerase chain reaction Methods 0.000 description 1
- 108091033319 polynucleotide Proteins 0.000 description 1
- 239000002157 polynucleotide Substances 0.000 description 1
- 102000040430 polynucleotide Human genes 0.000 description 1
- 239000002243 precursor Substances 0.000 description 1
- 108010045530 proricin Proteins 0.000 description 1
- 244000086221 rape kale Species 0.000 description 1
- 230000006798 recombination Effects 0.000 description 1
- 238000005215 recombination Methods 0.000 description 1
- 230000002829 reductive effect Effects 0.000 description 1
- 230000001172 regenerating effect Effects 0.000 description 1
- 230000008929 regeneration Effects 0.000 description 1
- 238000011069 regeneration method Methods 0.000 description 1
- 230000001850 reproductive effect Effects 0.000 description 1
- 150000008223 ribosides Chemical class 0.000 description 1
- 210000003705 ribosome Anatomy 0.000 description 1
- MEFOUWRMVYJCQC-UHFFFAOYSA-N rimsulfuron Chemical compound CCS(=O)(=O)C1=CC=CN=C1S(=O)(=O)NC(=O)NC1=NC(OC)=CC(OC)=N1 MEFOUWRMVYJCQC-UHFFFAOYSA-N 0.000 description 1
- HBMJWWWQQXIZIP-UHFFFAOYSA-N silicon carbide Chemical compound [Si+]#[C-] HBMJWWWQQXIZIP-UHFFFAOYSA-N 0.000 description 1
- 229910010271 silicon carbide Inorganic materials 0.000 description 1
- 239000001509 sodium citrate Substances 0.000 description 1
- NLJMYIDDQXHKNR-UHFFFAOYSA-K sodium citrate Chemical compound O.O.[Na+].[Na+].[Na+].[O-]C(=O)CC(O)(CC([O-])=O)C([O-])=O NLJMYIDDQXHKNR-UHFFFAOYSA-K 0.000 description 1
- 238000004162 soil erosion Methods 0.000 description 1
- 239000007787 solid Substances 0.000 description 1
- 239000002904 solvent Substances 0.000 description 1
- 230000000392 somatic effect Effects 0.000 description 1
- 238000005507 spraying Methods 0.000 description 1
- 239000008223 sterile water Substances 0.000 description 1
- 150000003431 steroids Chemical class 0.000 description 1
- 238000003756 stirring Methods 0.000 description 1
- 238000003860 storage Methods 0.000 description 1
- 239000000758 substrate Substances 0.000 description 1
- 239000006228 supernatant Substances 0.000 description 1
- 239000004094 surface-active agent Substances 0.000 description 1
- 230000004083 survival effect Effects 0.000 description 1
- 238000004114 suspension culture Methods 0.000 description 1
- RYYWUUFWQRZTIU-UHFFFAOYSA-K thiophosphate Chemical compound [O-]P([O-])([O-])=S RYYWUUFWQRZTIU-UHFFFAOYSA-K 0.000 description 1
- 238000003971 tillage Methods 0.000 description 1
- 239000003053 toxin Substances 0.000 description 1
- 231100000765 toxin Toxicity 0.000 description 1
- 229940035893 uracil Drugs 0.000 description 1
- 230000002792 vascular Effects 0.000 description 1
- 235000013311 vegetables Nutrition 0.000 description 1
- 238000003260 vortexing Methods 0.000 description 1
Classifications
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/11—DNA or RNA fragments; Modified forms thereof; Non-coding nucleic acids having a biological activity
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
- C12N15/8274—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for herbicide resistance
-
- C—CHEMISTRY; METALLURGY
- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
- C07K14/415—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from plants
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/10—Transferases (2.)
- C12N9/1022—Transferases (2.) transferring aldehyde or ketonic groups (2.2)
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
- C12N9/88—Lyases (4.)
Landscapes
- Health & Medical Sciences (AREA)
- Life Sciences & Earth Sciences (AREA)
- Genetics & Genomics (AREA)
- Chemical & Material Sciences (AREA)
- Engineering & Computer Science (AREA)
- Organic Chemistry (AREA)
- Wood Science & Technology (AREA)
- Zoology (AREA)
- Bioinformatics & Cheminformatics (AREA)
- Biomedical Technology (AREA)
- Biotechnology (AREA)
- Molecular Biology (AREA)
- General Engineering & Computer Science (AREA)
- Biochemistry (AREA)
- General Health & Medical Sciences (AREA)
- Microbiology (AREA)
- Biophysics (AREA)
- Medicinal Chemistry (AREA)
- Physics & Mathematics (AREA)
- Plant Pathology (AREA)
- Cell Biology (AREA)
- Botany (AREA)
- Gastroenterology & Hepatology (AREA)
- Proteomics, Peptides & Aminoacids (AREA)
- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
- Micro-Organisms Or Cultivation Processes Thereof (AREA)
- Agricultural Chemicals And Associated Chemicals (AREA)
- Enzymes And Modification Thereof (AREA)
- Preparation Of Compounds By Using Micro-Organisms (AREA)
- Catching Or Destruction (AREA)
- Measuring Or Testing Involving Enzymes Or Micro-Organisms (AREA)
- Plural Heterocyclic Compounds (AREA)
Description
本出願は,米国特許仮出願60/977,944(2007年10月5日出願)に基づく優先権を主張しており,この出願は,明細書,図面,表を含めその全体がすべての目的のために参照として本明細書に組み込まれる。
本発明は,除草剤耐性植物および種子の分野に関し,特に,アセトヒドロキシ酸シンターゼ(AHAS)遺伝子および蛋白質における変異に関する。
以下の説明は,単に本発明の理解を助けるために提供され,本発明に対する従来技術を記載するかまたは構成すると認めるものではない。
本発明は,部分的には,変異アセトヒドロキシ酸シンターゼ(AHAS)核酸,および変異した核酸によりコードされる蛋白質に関する。本発明はまた,部分的には,これらの変異した核酸および蛋白質を含むキャノーラ植物,細胞,および種子に関する。
本発明は,部分的には,例えば,サイバス社により開発された"the Rapid Trait Development System(RTDS(登録商標)"技術を用いて,アブラナ属のアセトヒドロキシ酸シンターゼ(AHAS)遺伝子を首尾良く標的化することに関連する組成物および方法を提供する。本明細書に開示されるいすれかの変異を含む植物は,組み合わせでまたは単独で,新規な除草剤耐性製品の基礎を形成することができる。また,AHAS遺伝子の変異についてホモ接合性またはヘテロ接合性のいずれかである変異した植物から生成される種子も提供される。本明細書に開示される変異は,他の既知の任意の変異との,または将来発見される変異との組み合わせであってもよい。
本明細書に開示される方法および組成物は,下記に詳細に説明されるコンフォメーションおよび化学を有する"遺伝子修復オリゴ核酸塩基"を用いて実施または製造することができる。本明細書において企図される"遺伝子修復オリゴ核酸塩基"とは,公開された化学文献および特許文献において,例えば,"組換え誘導性(recombinagenic)オリゴ核酸塩基";"RNA/DNAキメラオリゴヌクレオチド";"キメラオリゴヌクレオチド";"混合デュープレックスオリゴヌクレオチド"(MDONs);"RNADNAオリゴヌクレオチド(RDOs)";"遺伝子標的化オリゴヌクレオチド;""ジェノプラスト";"一本鎖修飾オリゴヌクレオチド";"一本鎖オリゴデオキシヌクレオチド変異用ベクター"(SSOMVs);"デュープレックス変異用ベクター";および"ヘテロデュープレックス変異用ベクター"等の他の名称を用いて記載されている。
植物細胞をトランスフォームするのに用いられる任意の既知の方法を用いて,遺伝子修復オリゴ核酸塩基をデリバリーすることができる。例示的な方法について以下に説明する。
セルロース細胞壁を有する植物細胞中に大きなフラグメントのDNAを発射貫入により導入するための金属マイクロキャリア(微小球)の使用は,当業者によく知られている(以下,バイオリスティックデリバリーと称する)。米国特許4,945,050;5,100,792および5,204,253は,マイクロキャリアおよびこれらを発射させるためのデバイスを選択する一般的手法を記載する。
別の態様においては,遺伝子修復オリゴ核酸塩基は,植物の一部に由来するプロトプラストのエレクトロポレーションにより植物細胞にデリバリーすることができる。プロトプラストは,当業者によく知られる手法にしたがって,植物の一部,特に葉を酵素で処理することにより形成される。例えば,Gallois et al.,1996,Methods in Molecular Biology 55:89−107,Humana Press,Totowa,N.J.;Kipp et al.,1999,Methods in Molecular Biology 133:213−221,Humana Press,Totowa,N.Jを参照。プロトプラストは,エレクトロポレーション前に成長培地で培養する必要はない。エレクトロポレーションの条件の一例は,総容量0.3mL中に3x105個のプロトプラストであり,遺伝子修復オリゴ核酸塩基の濃度は0.6−4μg/mLである。
別の態様においては,当業者によく知られる手法にしたがって膜改変剤であるポリエチレングリコールの存在下で,核酸を植物プロトプラストに取り込ませる(例えば,Gharti−Chhetri et al.,1992;Datta et al.,1992を参照)。
別の態様においては,遺伝子修復オリゴ核酸塩基は,マイクロキャピラリーを用いて植物細胞またはプロトプラストに注入することによりデリバリーすることができる(例えば,Miki et al.,1989;Schnorf et al.,1991を参照)。
植物および植物細胞は,当該技術分野において一般に知られる方法を用いて,例えば,植物または植物細胞を除草剤の存在下で生長させ,除草剤の非存在下における生長速度と比較した生長の速度を測定することにより,除草剤に対する耐性または寛容性について試験することができる。
本明細書において用いる場合,特に記載しないかぎり,遺伝子の番号付けは,シロイナズナのアセト乳酸シンターゼ(ALS)またはアセトヒドロキシ酸シンターゼ(AHAS)At3g48560(配列番号1)のアミノ酸配列に基づく。2005年10月より前の実験ノートでは,S653位(シロイナズナアミノ酸配列に基づく)は,トウモロコシのアミノ酸配列ZmAHAS108およびZmAHAS109(Fang et al.,1992)に基づいてS621として示されている。
細胞培養の説明
種子から,および小胞子から誘導された胚からの両方の新芽をインビトロで無菌条件下で増殖させた。2−4週間ごとに切片を継代し,ペトリ皿(25mmx90mm)で40−45mLの容量のRS培地(Dovzhenko,2001)で培養した。皿をミクロポアテープ(Microporetape,3M Company)で密封した。若葉をプロトプラスト単離に用いた。
インビトロで発芽した2−3週齢の葉組織約600mgを,5mLの培地B(Pelletier et al.,1983,pHを5.8に調節)を入れたペトリ皿中で,外科用メスを用いて小さいストリップに切断した。約1時間後,培地Bを,0.5%(w/v)のセルラーゼ(Cellulase)YCおよび0.75%(w/v)のマセロザイム(Macerozyme)R10(いずれもKarlan Research Products,Cottonwood,Arizonaより),1g/Lのウシ血清アルブミン,および1g/Lの2−モルホリノエタンスルホン酸を溶解した培地Bからなる酵素溶液で置き換えた。酵素溶液を葉組織中に減圧浸潤させ,酵素溶液中の葉切片を入れた皿を暗所で25℃でインキュベーションした。プロトプラスト精製は,イオジキサノール密度勾配(Optiprep Application Sheet C18;Purification of Intact Plant Protoplasts;Axis−Shield USA,10 Commerce Way,Norton,MA02776を適用)を用いて行った。密度勾配遠心分離の後,精製されたプロトプラストを含むバンドを約5mLのW5培地とともに取り出した(Frigerio et al.,1998)。プロトプラストの収率は血球計で測定し,プロトプラストは4℃で2時間保存した。
プロトプラスト懸濁液を等量のW5培地と混合し,50mLの遠心管に移し,臨床用遠心分離器で最低の設定(約50xg)で5分間遠心分離した。上清を除去し,TM培地(Klaus,2001)で置き換えて,プロトプラスト密度を5x106/mLに調節した。それぞれ5x106個のプロトプラストを含む100μLのアリコートを12mLの丸底遠心管に入れた。次に,AHAS遺伝子中の一方または両方の変異を標的とするGRONを,PEG処理を用いてプロトプラストに導入した。GRONをプロトプラストに導入するためには,12.5μgのGRONを25μLの純粋に溶解し,125μLのポリエチレングリコール溶液(5gPEG,MW1500,638mgマンニトール,207mgCaNO3x4H2Oおよび8.75mL純水;pHを約9.0に調節)を加えた。氷上で30分間インキュベーションした後,プロトプラスト−PEG懸濁液をW5培地で洗浄し,培地Bに再懸濁した。懸濁液を冷蔵庫で約4℃で一晩保持した。
GRONを導入した1日後,プロトプラストをアルギン酸カルシウムに包埋した。プロトプラストをゲル基質(例えば,アガロース,アルギン酸)中に包埋すると,プロトプラストの生存が高まり,プロトプラストに由来する細胞の***頻度が増加することが示されている。適用した方法は,Dovzhenko(2001)に記載されている方法に基づくものである。
イマゼタピル耐性カルスの選択は,Pelletier et al.(1983)にしたがって,培地中のアルギン酸の一連の継代培養により行った。0.5μMイマゼタピルの濃度でPEG/GRON処理した1週間後に選択を開始した。除草剤は即時効果は有していなかった。イマゼタピルなしで培養初期段階に形成されたすべてのマイクロコロニーは,最初は生長し続けたが,除草剤を加えていない対照より遅かった。選択開始の1から2週間後,コロニーの生長は鈍化するかまたは停止した。
固化選択培地上に発生し,そのDNAを分析すると変異の存在を示すImi寛容性カルスを,除草剤を含まない培地E(Pelletier et al.,1983)に移して,発生を加速させた。個々のカルス系統は,その生長速度および形態学は様々であった。一般に,新芽生成に向けた発生は以下の段階を経た:
未分化,緑色カルス−>暗緑色の領域をもつカルス−>発根−>始原細胞発芽−>過水状態の(ガラス化した)葉をもつ発育不良の発芽
5−6葉期のセイヨウアブラナ植物に種々のAHAS阻害除草剤を散布した。散布したB.napus植物には,親系統BN02(または必要な場合にはBN11),BN15(クリアフィールド(Clearfield),市販の2遺伝子チェック)および下記に示す変異体が含まれる。除草剤は,0.25%のAU391界面活性剤の存在下で下記の割合で散布した。
イマザモックス (Beyond(登録商標)) 0, 2, 4, 6, 8, 12, 16, 32 および 48 オンスの活性成分/エーカー(ai/A)
チフェンスルフロン 0.028, 0.056, 0.112, 0.168 ポンドの ai/A
トリベヌロン 0.015, 0.03, 0.06, 0.12, 0.18 ポンドのai/A
ニコスルフロン 0.06, 0.120, 0.24, 0.36 ポンドのai/A
リムスルフロン 0.015, 0.03, 0.06, 0.12, 0.18 ポンドのai/A
2:1 重量:重量チフェンスルフロン/トリベヌロン 0.056, 0.112, 0.224, 0.336 ポンドのai/A
2.22:1 チフェンスルフロン/ニコスルフロン 0.058, 0.116, 0.232 ポンドのai/A
プリミスルフロン 0.035, 0.070, 0.140 ポンドのai/A
フルメツラム0.040, 0.080, 0.16 ポンドのai/A
クロランスラム0.039, 0.078, 0.156 ポンドのai/A
チフェンスルフロン: BN2 (6), BN15 (6), BN15xBnALS-57 (18)
トリベヌロン: BN2 (6), BN15 (6), BN15xBnALS-57 (18), 63 (9)
ニコスルフロン: BN2 (6), BN15 (6), BN15xBnALS-57 (18)
THI / TRI: BN2 (6), BN15 (6), BN15xBnALS-57 (18)
リムスルフロン: BN2 (6), BN15 (6), BN15xBnALS-57 (18), 63 (9)
THI / Nic: BN2 (18), 63 (9), BN15xBnALS-57 (36), BN15 (18)
プリミスルフロン: BN2 (9), 63 (9), BN15 (9), BN15xBnALS-57 (18)
フルメツラム: BN2 (9), 63 (9), BN15 (9), BN15xBnALS-57 (18)
クロランスラム: BN2 (9), 63 (9), BN15 (9), BN15xBnALS-57 (18)
10回目の評点: 8, 16, 32, および 48 オンス/エーカー(oz/A)の BnALS-83xBnALS-123, BnALS-96xBnALS-123, BnALS-97xBnALS-57, BN15xBnALS-57, BN2, BN15, BnALS-97xBN15
17回目の評点: 4 および 12 oz/A の BnALS-97xBnALS-57, BN15xBnALS-97, BN2
28回目の評点: すべての単一因子変異について2, 4, 6, 8 oz/A
イマザモックス:
2 oz/A: BN2 (18), BnALS-123 (9), BnALS-96 (9), BnALS-97 (18), BnALS-83 (6), BnALS-76 (18), BnALS-58 (9), BnALS-57 (12)
4 oz/A: BN2 (18), BnALS-123 (9), BnALS-96 (9), BnALS-97 (18), BnALS-83 (9), BnALS-76 (18), BnALS-58 (9), BnALS-57 (12), BnALS-97xBN15 (15), BnALS-97xBnALS-57 (14)
6 oz/A: BN2 (9), BnALS-123 (9), BnALS-96 (12), BnALS-97 (9), BnALS-83 (12), BnALS-76 (9), BnALS-57 (12)
8 oz/A: BnALS-123 (9), BnALS-96 (12), BnALS-83 (12), BnALS-83xBnALS-123 (18), BnALS-96xBnALS-123 (18), BN2 (18), BN15 (18), BN15xBnALS-57 (15), BnALS-97xBnALS-57 (18), BN15xBnALS-97 (18)
12 oz/A: BnALS-97xBN15 (15), BnALS-97xBnALS-57 (14), BN2 (12)
16 oz/A: BnALS-83xBnALS-123 (18), BnALS-96xBnALS-123 (18), BN2 (15), BN15 (18), BN15xBnALS-57 (18), BnALS-97xBnALS-57 (18), BN15xBnALS-97 (18)
32 oz/A: BnALS-83xBnALS-123 (18), BnALS-96xBnALS-123 (18), BN2 (13), BN15 (18), BN15xBnALS-57 (18), BnALS-97xBnALS-57 (18), BN15xBnALS-97 (18)
48 oz/A: BnALS-83xBnALS-123 (18), BnALS-96xBnALS-123 (18), BN2 (6), BN15 (18), BN15xBnALS-57 (18), BnALS-97xBnALS-57 (18), BN15xBnALS-97 (18)
Datta SK, Datta K, Soltanifar N, Donn G, Potrykus I (1992) Herbicide-resistant Indica rice plants from IRRI breeding line IR72 after PEG-mediated transformation of protoplasts. Plant Molec. Biol. 20:619-629
Dovzhenko A (2001) Towards plastid transformation in rapeseed (Brassica napus L.) and sugarbeet (Beta vulgaris L.). PhD Dissertation, LMU Munich, Faculty of Biology
Edwards K, Johnstone C, Thompson C (1991) A simple and rapid method for the preparation of plant genomic DNA for PCR analysis. Nucleic Acids Res. 19:1349.
Frigerio L, Vitale A, Lord JM, Ceriotti A, Roberts LM (1998) Free ricin A chain, proricin, and native toxin have different cellular fates when expressed in tobacco protoplasts. J Biol Chem 273:14194-14199
Fang LY, Gross PR, Chen CH, Lillis M (1992) Sequence of two acetohydroxyacid synthase genes from Zea mays. Plant Mol Biol. 18(6):1185-7
Gharti-Chhetri GB, Cherdshewasart W, Dewulf J, Jacobs M, Negrutiu I (1992) Polyethylene glycol-mediated direct gene transfer in Nicotiana spp. Physiol. Plant. 85:345-351
Klaus S (2003) Markerfreie transplastome Tabakpflanzen (Marker-free transplastomic tobacco plants). PhD Dissertation, LMU Munich, Faculty of Biology
Miki B, Huang B, Bird S, Kemble R, Simmonds D, Keller W (1989) A procedure for the microinjection of plant cells and protoplasts. Meth. Cell Science 12:139-144
Pelletier G, Primard C, Vedel F, Chetrit P, Remy R, Rouselle P, Renard M (1983) Intergeneric cytoplasm hybridization in Cruciferae by protoplast fusion. Mol. Gen. Genet. 191: 244-250
Schnorf M, Neuhaus-Url G, Galli A, Iida S, Potrykus I, Neuhaus G (1991) An improved approach for transformation of plant cells by microinjection: molecular and genetic analysis. Transgen. Res. 1:23- 30
Tan S, Evans RR, Dahmer ML, Singh BK, Shaner DL (2005) Imidazolinone-tolerant crops: history, current status and future. Pest Manag Sci. 61(3):246-57
Claims (12)
- 除草剤耐性植物を製造する方法であって,
(a) 植物細胞に,アセトヒドロキシ酸シンターゼ(AHAS)I遺伝子を標的とした
変異を有する遺伝子修復オリゴ核酸塩基(GRON)を導入して,
配列番号1の574位に対応する位置におけるトリプトファンからシステインへの置換;
配列番号1の574位に対応する位置におけるトリプトファンからロイシンへの置換,
配列番号1の574位に対応する位置におけるトリプトファンからメチオニンへの置換;および
配列番号1の574位に対応する位置におけるトリプトファンからセリンへの置換;
からなる群より選択される変異を含むAHAS蛋白質を発現するAHAS遺伝子Iを有す
る植物細胞を生成し;
(b) AHAS阻害性除草剤の存在下で,対応する野生型植物細胞と比較して実質的に正常な生長および触媒活性を有する植物細胞を同定し;そして
(c) 前記植物細胞から変異したAHAS遺伝子Iを有する非トランスジェニック除草
剤耐性植物を再生する,
の各工程を含む方法。 - 変異は配列番号1の574位に対応する位置におけるトリプトファンからロイシンへの置換を含む,請求項1に記載の方法。
- 前記AHAS遺伝子IはAHAS阻害性除草剤による阻害に対して耐性である蛋白質をコ
ードする,請求項1記載の方法。 - 前記AHAS阻害性除草剤は,イミダゾリノン,スルホニルウレア,トリアゾロピリミジン,ピリミジニルチオベンゾエート,スルホニルアミノ−カルボニルトリアゾリノン,およびこれらの混合物からなる群より選択される除草剤である,請求項3に記載の方法。
- 前記AHAS阻害性除草剤はイミダゾリノン除草剤である,請求項4に記載の方法。
- 前記AHAS阻害性除草剤はスルホニルウレア除草剤である,請求項4に記載の方法。
- 前記AHAS遺伝子Iは,配列番号21〜62のアミノ酸配列の1またはそれ以上と90
%またはそれ以上の同一性を含む蛋白質をコードする,請求項1〜6のいずれか一項に記載の方法。 - 前記植物は,キャノーラ,ヒマワリ,タバコ,テンサイ,綿花,トウモロコシ,コムギ,オオムギ,イネ,ソルガム,トマト,マンゴー,モモ,リンゴ,洋ナシ,イチゴ,バナナ,メロン,ジャガイモ,ニンジン,レタス,タマネギ,大豆種,サトウキビ,エンドウ,ソラマメ,ポプラ,ブドウ,柑橘類,アルファルファ,ライ麦,オートムギ,芝草および飼料草,亜麻,菜種,キュウリ,アサガオ,ニガウリ,ピーマン,ナス,マリーゴールド,ハス,キャベツ,デイジー,カーネーション,チューリップ,アヤメ,およびユリからなる群の植物の種から選択される、請求項1〜7のいずれか一項に記載の方法。
- 前記植物はアブラナ属である、請求項1〜8のいずれか一項に記載の方法。
- 除草剤耐性植物を製造する方法であって,
(a) アセトヒドロキシ酸シンターゼ(AHAS)遺伝子Iを標的とした変異を有する
遺伝子修復オリゴ核酸塩基(GRON)を植物細胞に導入して,
配列番号1の574位に対応する位置におけるトリプトファンからシステインへの置換;
配列番号1の574位に対応する位置におけるトリプトファンからロイシンへの置換,
配列番号1の574位に対応する位置におけるトリプトファンからメチオニンへの置換;および
配列番号1の574位に対応する位置におけるトリプトファンからセリンへの置換,
からなる群より選択される変異を含むAHAS蛋白質を発現するAHAS遺伝子Iを有す
る植物細胞を生成し;
(b) AHAS阻害性除草剤の存在下で対応する野生型植物細胞と比較して実質的に正常な生長および触媒活性を有する植物細胞を同定し;そして
(c) 前記植物細胞から変異したAHAS遺伝子Iを有する非トランスジェニック除草
剤耐性植物を再生する,
の各工程を含み,
ここで,前記除草剤耐性植物はテンサイであり,前記除草剤はスルホニルウレア除草剤である,ことを特徴とする方法。 - 変異は配列番号1の574位に対応する位置におけるトリプトファンからロイシンへの置換を含む,請求項10に記載の方法。
- 前記AHAS遺伝子Iは,配列番号21〜62のアミノ酸配列の1またはそれ以上と90
%またはそれ以上の同一性を含む蛋白質をコードする,請求項10または11に記載の方法。
Applications Claiming Priority (3)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
US97794407P | 2007-10-05 | 2007-10-05 | |
US60/977,944 | 2007-10-05 | ||
PCT/US2008/078798 WO2009046334A1 (en) | 2007-10-05 | 2008-10-03 | Mutated acetohydroxyacid synthase genes in brassica |
Related Child Applications (1)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
JP2015094818A Division JP6092930B2 (ja) | 2007-10-05 | 2015-05-07 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
Publications (3)
Publication Number | Publication Date |
---|---|
JP2010539986A JP2010539986A (ja) | 2010-12-24 |
JP2010539986A5 JP2010539986A5 (ja) | 2011-11-17 |
JP5745849B2 true JP5745849B2 (ja) | 2015-07-08 |
Family
ID=40526700
Family Applications (4)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
JP2010528175A Active JP5745849B2 (ja) | 2007-10-05 | 2008-10-03 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
JP2015094818A Active JP6092930B2 (ja) | 2007-10-05 | 2015-05-07 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
JP2017022074A Active JP6529528B2 (ja) | 2007-10-05 | 2017-02-09 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
JP2019091122A Active JP6833903B2 (ja) | 2007-10-05 | 2019-05-14 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
Family Applications After (3)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
JP2015094818A Active JP6092930B2 (ja) | 2007-10-05 | 2015-05-07 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
JP2017022074A Active JP6529528B2 (ja) | 2007-10-05 | 2017-02-09 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
JP2019091122A Active JP6833903B2 (ja) | 2007-10-05 | 2019-05-14 | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 |
Country Status (13)
Country | Link |
---|---|
US (3) | US20090205064A1 (ja) |
EP (4) | EP3584325A1 (ja) |
JP (4) | JP5745849B2 (ja) |
CN (5) | CN102405846A (ja) |
AU (1) | AU2008308530A1 (ja) |
BR (3) | BRPI0818295B1 (ja) |
CA (1) | CA2701624C (ja) |
DK (3) | DK2700721T3 (ja) |
EA (3) | EA036997B1 (ja) |
LT (2) | LT2700721T (ja) |
PL (3) | PL2700722T3 (ja) |
UA (2) | UA119225C2 (ja) |
WO (1) | WO2009046334A1 (ja) |
Families Citing this family (42)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
UA108733C2 (uk) | 2006-12-12 | 2015-06-10 | Толерантна до гербіциду рослина соняшника | |
NZ597328A (en) * | 2007-04-04 | 2014-03-28 | Basf Se | Herbicide-resistant brassica plants and methods of use |
US10017827B2 (en) | 2007-04-04 | 2018-07-10 | Nidera S.A. | Herbicide-resistant sunflower plants with multiple herbicide resistant alleles of AHASL1 and methods of use |
KR101452818B1 (ko) * | 2007-12-21 | 2014-10-23 | 키진 엔.브이. | 식물 원형질체 내로 폴리에틸렌 글리콜 매개 돌연변이 뉴클레오염기의 도입을 이용한 개선된 돌연변이 생성방법 |
AU2013200838B2 (en) * | 2007-12-21 | 2014-02-27 | Keygene N.V. | An improved mutagenesis method using polyethylene glycol mediated introduction of mutagenic nucleobases into plant protoplasts |
US8889951B2 (en) * | 2008-05-06 | 2014-11-18 | Agriculture Victoria Services Pty Ltd | Herbicide resistant barley expressing mutant acetohydroxy acid synthase |
CN102216453B (zh) * | 2008-09-26 | 2014-02-05 | 巴斯夫农化产品有限公司 | 除草剂-抗性的ahas-突变体及使用方法 |
CN102666859A (zh) | 2009-12-22 | 2012-09-12 | 拜尔作物科学公司 | 耐除草剂的植物 |
AU2011204287A1 (en) | 2010-01-07 | 2012-08-30 | BASF Agro B.V. | Herbicide-tolerant plants |
DK2547766T3 (en) | 2010-03-17 | 2018-04-16 | Basf Agrochemical Products Bv | Herbicide tolerant plants |
UA108751C2 (uk) * | 2010-03-17 | 2015-06-10 | Спосіб боротьби з бур'янами в озимій культурі brassica | |
UA112969C2 (uk) * | 2010-08-03 | 2016-11-25 | Сібас Юс Ллс | Рослина, стійка до одного або більше ррх-інгібуючих гербіцидів, яка містить мутантний ген протопорфіриноген ix оксидази (ррх) |
US20120122223A1 (en) | 2010-08-03 | 2012-05-17 | Cibus Us Llc | Mutated protoporphyrinogen ix oxidase (ppx) genes |
PT2627183T (pt) | 2010-10-15 | 2017-12-11 | Bayer Ip Gmbh | Utilização de herbicidas inibidores de als para controlo de vegetação indesejada em plantas beta vulgaris tolerantes a inibidor de als |
ES2954135T3 (es) | 2010-10-15 | 2023-11-20 | Bayer Cropscience Lp | Mutantes de Beta vulgaris tolerantes a herbicidas inhibidores de ALS |
UA117447C2 (uk) | 2011-05-04 | 2018-08-10 | Байєр Інтеллектуал Проперті Гмбх | Застосування гербіцидів інгібіторів als для боротьби з небажаною рослинністю на площі вирощування brassica |
US9572315B2 (en) * | 2011-05-04 | 2017-02-21 | Bayer Intellectual Property Gmbh | ALS inhibitor herbicide tolerant B. napus mutants |
CA2865571A1 (en) | 2012-02-29 | 2013-09-06 | Bayer Cropscience Nv | Als inhibitor herbicide tolerant b. napus mutants |
EP2834361A1 (en) * | 2012-04-05 | 2015-02-11 | Advanta International BV | Sorghum plants having a mutant polynucleotide encoding the large subunit of mutated acetohydroxyacid synthase protein and increased resistance to herbicides |
LT2931034T (lt) * | 2012-12-13 | 2018-10-10 | Bayer Cropscience Ag | Als slopinančių herbicidų panaudojimas nepageidaujamos augmenijos kontrolei augaluose beta vulgaris, kurie tolerantiški als slopinantiems herbicidams |
EP2931902B1 (en) * | 2012-12-13 | 2018-08-01 | SESVanderHave N.V. | Method to develop herbicide-resistant sugar beet plants |
AR095109A1 (es) | 2013-01-31 | 2015-09-30 | Univ Guelph | Plantas tolerantes a herbicidas auxínicos |
US20230313211A1 (en) * | 2013-03-15 | 2023-10-05 | Cibus Us Llc | Methods and compositions for increasing efficiency of targeted gene modification using oligonucleotide-mediated gene repair |
US9957515B2 (en) | 2013-03-15 | 2018-05-01 | Cibus Us Llc | Methods and compositions for targeted gene modification |
HUE058351T2 (hu) | 2013-03-15 | 2022-07-28 | Cibus Us Llc | Célzott génmódosítás oligonukleotid-közvetített génjavítás alkalmazásával |
RS63427B1 (sr) * | 2013-03-15 | 2022-08-31 | Cibus Us Llc | Postupci i kompozicije za povećanje efikasnosti modifikacije ciljnog gena korišćenjem genske reparacije posredovane oligonukleotidom |
CN103215243B (zh) * | 2013-04-07 | 2014-10-08 | 深圳兴旺生物种业有限公司 | 甘蓝型油菜抗除草剂蛋白及其在植物育种中的应用 |
EP3019617A1 (en) * | 2013-07-12 | 2016-05-18 | Bayer CropScience NV | Als inhibitor herbicide tolerant mutant plants |
US9072298B2 (en) | 2013-11-01 | 2015-07-07 | Rotam Agrochem Intrnational Company Limited | Synergistic herbicidal composition |
WO2015082413A1 (en) * | 2013-12-02 | 2015-06-11 | Bayer Cropscience Nv | Als inhibitor herbicide tolerant mutant plants |
KR102450868B1 (ko) * | 2014-03-14 | 2022-10-06 | 시버스 유에스 엘엘씨 | 올리고뉴클레오타이드 매개 유전자 보수를 사용한 표적화된 유전자 변형의 효율을 증가시키기 위한 방법 및 조성물 |
US20160138040A1 (en) * | 2014-11-13 | 2016-05-19 | Cellectis | Brassica engineered to confer herbicide tolerance |
WO2016130247A1 (en) | 2015-01-10 | 2016-08-18 | Cibus Us Llc | Mutated acetohydroxyacid synthase genes in euphorbiaceae and plant material comprising such genes |
CN104789682B (zh) * | 2015-04-29 | 2019-05-17 | 江苏省农业科学院 | 检测甘蓝型油菜抗磺酰脲类除草剂基因BnALS3R的引物与应用 |
CA3014036A1 (en) * | 2016-02-09 | 2017-08-17 | Cibus Us Llc | Methods and compositions for increasing efficiency of targeted gene modification using oligonucleotide-mediated gene repair |
CN107058350A (zh) * | 2017-03-20 | 2017-08-18 | 江苏省农业科学院 | 基于体外定点突变获得的油菜抗多种als抑制剂类除草剂基因及应用 |
US11021697B2 (en) | 2017-07-11 | 2021-06-01 | Cj Cheiljedang Corporation | Acetohydroxy acid synthase variant, microorganism comprising the same, and method of producing L-branched-chain amino acid using the same |
KR101996129B1 (ko) | 2017-07-11 | 2019-07-04 | 씨제이제일제당 (주) | 아세토하이드록시산 신타아제 변이체, 이를 포함하는 미생물 또는 이를 이용하는 l-분지쇄 아미노산 생산 방법 |
CN109197576A (zh) * | 2018-11-16 | 2019-01-15 | 贵州省油料研究所 | 一种甘蓝油菜隐性核不育系与百合杂交选育油菜的方法 |
CN109880927A (zh) * | 2019-03-20 | 2019-06-14 | 江苏省农业科学院 | 检测油菜BnALS1R基因的SNP标记引物及其应用 |
CN110628783A (zh) * | 2019-10-23 | 2019-12-31 | 江苏省农业科学院 | 一种非转基因抗除草剂油菜基因及其应用 |
KR20240105550A (ko) * | 2022-12-28 | 2024-07-08 | 씨제이제일제당 (주) | 신규한 아세토락테이트 합성효소 활성을 갖는 변이체 폴리펩티드 및 이를 이용한 l-글루탐산 생산 방법 |
Family Cites Families (39)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
US4545060A (en) | 1983-09-19 | 1985-10-01 | Northern Telecom Limited | Decision feedback adaptive equalizer acting on zero states following a non-zero state |
US5100792A (en) * | 1984-11-13 | 1992-03-31 | Cornell Research Foundation, Inc. | Method for transporting substances into living cells and tissues |
US4945050A (en) * | 1984-11-13 | 1990-07-31 | Cornell Research Foundation, Inc. | Method for transporting substances into living cells and tissues and apparatus therefor |
IL83348A (en) * | 1986-08-26 | 1995-12-08 | Du Pont | Nucleic acid fragment encoding herbicide resistant plant acetolactate synthase |
US5302523A (en) | 1989-06-21 | 1994-04-12 | Zeneca Limited | Transformation of plant cells |
US5204253A (en) * | 1990-05-29 | 1993-04-20 | E. I. Du Pont De Nemours And Company | Method and apparatus for introducing biological substances into living cells |
TW208716B (ja) * | 1990-12-27 | 1993-07-01 | American Cyanamid Co | |
DE4216134A1 (de) * | 1991-06-20 | 1992-12-24 | Europ Lab Molekularbiolog | Synthetische katalytische oligonukleotidstrukturen |
US5731180A (en) * | 1991-07-31 | 1998-03-24 | American Cyanamid Company | Imidazolinone resistant AHAS mutants |
IT230274Y1 (it) | 1993-06-11 | 1999-06-02 | Silc Spa | Pannolone assorbente sagomato per incontinenza |
ATE196311T1 (de) * | 1993-12-09 | 2000-09-15 | Univ Jefferson | Verbindungen und verfahren zur ortsspezifischen mutation in eukaryotischen zellen |
PT679657E (pt) | 1994-04-27 | 2003-11-28 | Novartis Ag | Nucleosidos e oligonucleotidos com grupos 2'-eter |
US5780296A (en) | 1995-01-17 | 1998-07-14 | Thomas Jefferson University | Compositions and methods to promote homologous recombination in eukaryotic cells and organisms |
US6576455B1 (en) * | 1995-04-20 | 2003-06-10 | Basf Corporation | Structure-based designed herbicide resistant products |
US5853973A (en) * | 1995-04-20 | 1998-12-29 | American Cyanamid Company | Structure based designed herbicide resistant products |
US5760012A (en) | 1996-05-01 | 1998-06-02 | Thomas Jefferson University | Methods and compounds for curing diseases caused by mutations |
US5731181A (en) | 1996-06-17 | 1998-03-24 | Thomas Jefferson University | Chimeric mutational vectors having non-natural nucleotides |
US5888983A (en) * | 1996-05-01 | 1999-03-30 | Thomas Jefferson University | Method and oligonucleobase compounds for curing diseases caused by mutations |
US5859348A (en) * | 1996-07-17 | 1999-01-12 | Board Of Trustees Operating Michigan State University | Imidazolinone and sulfonyl urea herbicide resistant sugar beet plants |
US6524613B1 (en) | 1997-04-30 | 2003-02-25 | Regents Of The University Of Minnesota | Hepatocellular chimeraplasty |
EP0979311A1 (en) | 1997-04-30 | 2000-02-16 | Of The University Of Minnesota Regents | $i(IN VIVO) USE OF RECOMBINAGENIC OLIGONUCLEOBASES TO CORRECT GENETIC LESIONS IN HEPATOCYTES |
GB9711015D0 (en) * | 1997-05-28 | 1997-07-23 | Zeneca Ltd | Improvements in or relating to organic compounds |
US7094606B2 (en) | 1997-08-05 | 2006-08-22 | Arntzen Charles J | Use of mixed duplex oligonucleotides to effect localized genetic changes in plants |
US6004804A (en) | 1998-05-12 | 1999-12-21 | Kimeragen, Inc. | Non-chimeric mutational vectors |
US6010907A (en) | 1998-05-12 | 2000-01-04 | Kimeragen, Inc. | Eukaryotic use of non-chimeric mutational vectors |
US6271360B1 (en) * | 1999-08-27 | 2001-08-07 | Valigen (Us), Inc. | Single-stranded oligodeoxynucleotide mutational vectors |
AR025996A1 (es) * | 1999-10-07 | 2002-12-26 | Valigen Us Inc | Plantas no transgenicas resistentes a los herbicidas. |
US6936467B2 (en) * | 2000-03-27 | 2005-08-30 | University Of Delaware | Targeted chromosomal genomic alterations with modified single stranded oligonucleotides |
AU6135801A (en) * | 2000-05-10 | 2001-11-20 | Univ Louisiana State | Resistance to acetohydroxyacid synthase-inhibiting herbicides |
US20030097692A1 (en) * | 2000-12-21 | 2003-05-22 | Georg Jander | Plants with imidazolinone-resistant ALS |
CN101565708B (zh) * | 2002-03-29 | 2011-09-21 | 组合化学工业株式会社 | 编码乙酰乳酸合酶的基因 |
US20060130172A1 (en) * | 2003-01-09 | 2006-06-15 | Cory Whaley | Gene encoding resistance to acetolactate synthase-inhibiting herbicides |
MXPA06002155A (es) * | 2003-08-29 | 2007-01-25 | Inst Nac De Technologia Agrope | Plantas de arroz que tienen una tolerancia incrementada a los herbicidas de imidazolinona. |
UA97344C2 (uk) * | 2004-07-30 | 2012-02-10 | Басф Агрокемікел Продактс Б.В. | Резистентні до імідазолінонових гербіцидів рослини соняшнику , полінуклеотиди, що кодують резистентні до гербіцидів великі субодиниці білків ацетогідроксикислотної синтази |
AR051690A1 (es) * | 2004-12-01 | 2007-01-31 | Basf Agrochemical Products Bv | Mutacion implicada en el aumento de la tolerancia a los herbicidas imidazolinona en las plantas |
UA99091C2 (ru) * | 2005-07-01 | 2012-07-25 | Басф Се | Резистентное к гербициду растение подсолнечника, полинуклеотид, кодирующий резистентную к гербициду большую субъединицу белка ацетогидроксикислотной синтазы |
US20070118920A1 (en) * | 2005-11-09 | 2007-05-24 | Basf Agrochemical Products B.V. | Herbicide-resistant sunflower plants, polynucleotides encoding herbicide-resistant acetohydroxyacid synthase large subunit proteins, and methods of use |
UA108733C2 (uk) | 2006-12-12 | 2015-06-10 | Толерантна до гербіциду рослина соняшника | |
BRPI0810077B1 (pt) * | 2007-04-04 | 2018-04-24 | Basf Plant Science Gmbh | Polipeptídeo da subunidade grande da acetohidroxiácido sintase, e, métodos para controlar ervas daninhas na adjacência de plantas cultivadas, para identificar ou selecionar uma célula de planta transformada, tecido de planta, planta ou parte da mesma, e para combater vegetação indesejada |
-
2008
- 2008-10-03 CN CN2011101339067A patent/CN102405846A/zh active Pending
- 2008-10-03 BR BRPI0818295-7A patent/BRPI0818295B1/pt active IP Right Grant
- 2008-10-03 LT LTEP13175389.9T patent/LT2700721T/lt unknown
- 2008-10-03 UA UAA201309872A patent/UA119225C2/uk unknown
- 2008-10-03 AU AU2008308530A patent/AU2008308530A1/en not_active Abandoned
- 2008-10-03 EA EA201691664A patent/EA036997B1/ru unknown
- 2008-10-03 EP EP19174921.7A patent/EP3584325A1/en active Pending
- 2008-10-03 US US12/245,610 patent/US20090205064A1/en not_active Abandoned
- 2008-10-03 BR BR122018070230-3A patent/BR122018070230B1/pt active IP Right Grant
- 2008-10-03 CN CN201810145878.2A patent/CN108342376A/zh active Pending
- 2008-10-03 EA EA201070430A patent/EA026107B1/ru not_active IP Right Cessation
- 2008-10-03 WO PCT/US2008/078798 patent/WO2009046334A1/en active Application Filing
- 2008-10-03 PL PL13175393T patent/PL2700722T3/pl unknown
- 2008-10-03 UA UAA201005275A patent/UA103887C2/ru unknown
- 2008-10-03 EP EP13175393.1A patent/EP2700722B1/en active Active
- 2008-10-03 DK DK13175389.9T patent/DK2700721T3/en active
- 2008-10-03 EP EP08836084.7A patent/EP2203565B1/en active Active
- 2008-10-03 BR BR122018070228-1A patent/BR122018070228B1/pt active IP Right Grant
- 2008-10-03 CN CN201810149486.3A patent/CN108130336A/zh active Pending
- 2008-10-03 CA CA2701624A patent/CA2701624C/en active Active
- 2008-10-03 CN CN2008801190843A patent/CN101883868A/zh active Pending
- 2008-10-03 EP EP13175389.9A patent/EP2700721B1/en active Active
- 2008-10-03 LT LTEP13175393.1T patent/LT2700722T/lt unknown
- 2008-10-03 PL PL13175389T patent/PL2700721T3/pl unknown
- 2008-10-03 DK DK13175393.1T patent/DK2700722T3/da active
- 2008-10-03 PL PL08836084T patent/PL2203565T3/pl unknown
- 2008-10-03 JP JP2010528175A patent/JP5745849B2/ja active Active
- 2008-10-03 EA EA202092479A patent/EA202092479A3/ru unknown
- 2008-10-03 DK DK08836084.7T patent/DK2203565T3/en active
- 2008-10-03 CN CN202210655806.9A patent/CN115044593A/zh active Pending
-
2012
- 2012-02-28 US US13/407,676 patent/US20120178628A1/en not_active Abandoned
-
2015
- 2015-05-07 JP JP2015094818A patent/JP6092930B2/ja active Active
-
2017
- 2017-02-09 JP JP2017022074A patent/JP6529528B2/ja active Active
-
2019
- 2019-05-14 JP JP2019091122A patent/JP6833903B2/ja active Active
- 2019-12-27 US US16/728,666 patent/US20200123563A1/en not_active Abandoned
Also Published As
Similar Documents
Publication | Publication Date | Title |
---|---|---|
JP6833903B2 (ja) | アブラナ属の変異アセトヒドロキシ酸シンターゼ遺伝子 | |
JP7354205B2 (ja) | 変異したプロトポルフィリノーゲンixオキシダーゼ(ppx)遺伝子 | |
US10676755B2 (en) | Mutated acetohydroxyacid synthase genes in euphorbiaceae and plant material comprising such genes | |
AU2017279602B2 (en) | Mutated acetohydroxyacid synthase genes in Brassica | |
AU2022231679A1 (en) | Mutated acetohydroxyacid synthase genes in Brassica | |
NZ735454A (en) | Mutated acetohydroxyacid synthase genes in brassica |
Legal Events
Date | Code | Title | Description |
---|---|---|---|
A521 | Request for written amendment filed |
Free format text: JAPANESE INTERMEDIATE CODE: A523 Effective date: 20110930 |
|
A621 | Written request for application examination |
Free format text: JAPANESE INTERMEDIATE CODE: A621 Effective date: 20110930 |
|
A711 | Notification of change in applicant |
Free format text: JAPANESE INTERMEDIATE CODE: A711 Effective date: 20111114 |
|
A521 | Request for written amendment filed |
Free format text: JAPANESE INTERMEDIATE CODE: A821 Effective date: 20111114 |
|
A131 | Notification of reasons for refusal |
Free format text: JAPANESE INTERMEDIATE CODE: A131 Effective date: 20130604 |
|
A601 | Written request for extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A601 Effective date: 20130829 |
|
A602 | Written permission of extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A602 Effective date: 20130905 |
|
A601 | Written request for extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A601 Effective date: 20130930 |
|
A602 | Written permission of extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A602 Effective date: 20131007 |
|
A601 | Written request for extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A601 Effective date: 20131029 |
|
A602 | Written permission of extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A602 Effective date: 20131106 |
|
A521 | Request for written amendment filed |
Free format text: JAPANESE INTERMEDIATE CODE: A523 Effective date: 20131204 |
|
A131 | Notification of reasons for refusal |
Free format text: JAPANESE INTERMEDIATE CODE: A131 Effective date: 20140722 |
|
A601 | Written request for extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A601 Effective date: 20141016 |
|
A602 | Written permission of extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A602 Effective date: 20141023 |
|
A601 | Written request for extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A601 Effective date: 20141117 |
|
A602 | Written permission of extension of time |
Free format text: JAPANESE INTERMEDIATE CODE: A602 Effective date: 20141125 |
|
A521 | Request for written amendment filed |
Free format text: JAPANESE INTERMEDIATE CODE: A523 Effective date: 20141222 |
|
TRDD | Decision of grant or rejection written | ||
A01 | Written decision to grant a patent or to grant a registration (utility model) |
Free format text: JAPANESE INTERMEDIATE CODE: A01 Effective date: 20150407 |
|
A61 | First payment of annual fees (during grant procedure) |
Free format text: JAPANESE INTERMEDIATE CODE: A61 Effective date: 20150507 |
|
R150 | Certificate of patent or registration of utility model |
Ref document number: 5745849 Country of ref document: JP Free format text: JAPANESE INTERMEDIATE CODE: R150 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
S111 | Request for change of ownership or part of ownership |
Free format text: JAPANESE INTERMEDIATE CODE: R313117 |
|
R350 | Written notification of registration of transfer |
Free format text: JAPANESE INTERMEDIATE CODE: R350 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |
|
R250 | Receipt of annual fees |
Free format text: JAPANESE INTERMEDIATE CODE: R250 |