US20030167539A1 - Genetically modified plants having modulated flower development - Google Patents
Genetically modified plants having modulated flower development Download PDFInfo
- Publication number
- US20030167539A1 US20030167539A1 US09/849,772 US84977201A US2003167539A1 US 20030167539 A1 US20030167539 A1 US 20030167539A1 US 84977201 A US84977201 A US 84977201A US 2003167539 A1 US2003167539 A1 US 2003167539A1
- Authority
- US
- United States
- Prior art keywords
- nucleic acid
- plant
- seq
- leu
- glu
- 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.)
- Abandoned
Links
- 230000008124 floral development Effects 0.000 title claims description 19
- 241000196324 Embryophyta Species 0.000 claims abstract description 247
- 108090000623 proteins and genes Proteins 0.000 claims abstract description 108
- 102000004169 proteins and genes Human genes 0.000 claims abstract description 37
- 230000020429 meristem development Effects 0.000 claims abstract description 26
- 150000007523 nucleic acids Chemical group 0.000 claims description 92
- 210000004027 cell Anatomy 0.000 claims description 89
- 102000039446 nucleic acids Human genes 0.000 claims description 53
- 108020004707 nucleic acids Proteins 0.000 claims description 53
- 239000013598 vector Substances 0.000 claims description 45
- 238000000034 method Methods 0.000 claims description 44
- 241000219194 Arabidopsis Species 0.000 claims description 42
- 108091028043 Nucleic acid sequence Proteins 0.000 claims description 39
- 238000009396 hybridization Methods 0.000 claims description 24
- 230000001939 inductive effect Effects 0.000 claims description 14
- 240000001436 Antirrhinum majus Species 0.000 claims description 11
- 230000001105 regulatory effect Effects 0.000 claims description 10
- 210000001938 protoplast Anatomy 0.000 claims description 9
- 239000003550 marker Substances 0.000 claims description 7
- 241001406921 Squamosa Species 0.000 claims description 6
- 239000000126 substance Substances 0.000 claims description 5
- 230000001747 exhibiting effect Effects 0.000 claims description 4
- 239000002773 nucleotide Substances 0.000 claims 13
- 125000003729 nucleotide group Chemical group 0.000 claims 13
- 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 claims 4
- 101710116069 Floricaula protein Proteins 0.000 claims 4
- 235000002637 Nicotiana tabacum Nutrition 0.000 abstract description 28
- 241000183024 Populus tremula Species 0.000 abstract description 26
- 238000004519 manufacturing process Methods 0.000 abstract description 5
- 241000208125 Nicotiana Species 0.000 abstract 1
- 230000009466 transformation Effects 0.000 description 29
- 244000061176 Nicotiana tabacum Species 0.000 description 27
- 230000014509 gene expression Effects 0.000 description 26
- 230000017260 vegetative to reproductive phase transition of meristem Effects 0.000 description 26
- 241000282326 Felis catus Species 0.000 description 23
- 239000013612 plasmid Substances 0.000 description 22
- 235000018102 proteins Nutrition 0.000 description 22
- 230000009261 transgenic effect Effects 0.000 description 20
- 108020004414 DNA Proteins 0.000 description 19
- 230000000694 effects Effects 0.000 description 18
- 241000894007 species Species 0.000 description 17
- 210000000056 organ Anatomy 0.000 description 16
- 238000011161 development Methods 0.000 description 13
- 230000018109 developmental process Effects 0.000 description 13
- 230000035772 mutation Effects 0.000 description 13
- 241000589158 Agrobacterium Species 0.000 description 12
- 108010008355 arginyl-glutamine Proteins 0.000 description 11
- YBAFDPFAUTYYRW-UHFFFAOYSA-N N-L-alpha-glutamyl-L-leucine Natural products CC(C)CC(C(O)=O)NC(=O)C(N)CCC(O)=O YBAFDPFAUTYYRW-UHFFFAOYSA-N 0.000 description 10
- 108010005233 alanylglutamic acid Proteins 0.000 description 10
- XKUKSGPZAADMRA-UHFFFAOYSA-N glycyl-glycyl-glycine Chemical compound NCC(=O)NCC(=O)NCC(O)=O XKUKSGPZAADMRA-UHFFFAOYSA-N 0.000 description 10
- 238000011069 regeneration method Methods 0.000 description 10
- 241000701489 Cauliflower mosaic virus Species 0.000 description 9
- PMGDADKJMCOXHX-UHFFFAOYSA-N L-Arginyl-L-glutamin-acetat Natural products NC(=N)NCCCC(N)C(=O)NC(CCC(N)=O)C(O)=O PMGDADKJMCOXHX-UHFFFAOYSA-N 0.000 description 9
- 239000002299 complementary DNA Substances 0.000 description 9
- 230000006698 induction Effects 0.000 description 9
- 230000008929 regeneration Effects 0.000 description 9
- GCYFUZJHAXJKKE-KKUMJFAQSA-N Glu-Arg-Tyr Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O GCYFUZJHAXJKKE-KKUMJFAQSA-N 0.000 description 8
- 108010009111 arginyl-glycyl-glutamic acid Proteins 0.000 description 8
- 108010069495 cysteinyltyrosine Proteins 0.000 description 8
- 230000035784 germination Effects 0.000 description 8
- 108010025306 histidylleucine Proteins 0.000 description 8
- NWVVKQZOVSTDBQ-CIUDSAMLSA-N Ala-Glu-Arg Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O NWVVKQZOVSTDBQ-CIUDSAMLSA-N 0.000 description 7
- 102000053602 DNA Human genes 0.000 description 7
- RCCDHXSRMWCOOY-GUBZILKMSA-N Glu-Arg-Gln Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(N)=O)C(O)=O RCCDHXSRMWCOOY-GUBZILKMSA-N 0.000 description 7
- 241000218922 Magnoliophyta Species 0.000 description 7
- 230000015572 biosynthetic process Effects 0.000 description 7
- 108010009298 lysylglutamic acid Proteins 0.000 description 7
- 229920001184 polypeptide Polymers 0.000 description 7
- 108090000765 processed proteins & peptides Proteins 0.000 description 7
- 102000004196 processed proteins & peptides Human genes 0.000 description 7
- 108010077112 prolyl-proline Proteins 0.000 description 7
- VCSABYLVNWQYQE-UHFFFAOYSA-N Ala-Lys-Lys Natural products NCCCCC(NC(=O)C(N)C)C(=O)NC(CCCCN)C(O)=O VCSABYLVNWQYQE-UHFFFAOYSA-N 0.000 description 6
- ITHMWNNUDPJJER-ULQDDVLXSA-N Arg-His-Tyr Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O ITHMWNNUDPJJER-ULQDDVLXSA-N 0.000 description 6
- AGVNTAUPLWIQEN-ZPFDUUQYSA-N Arg-Ile-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N AGVNTAUPLWIQEN-ZPFDUUQYSA-N 0.000 description 6
- GHODABZPVZMWCE-FXQIFTODSA-N Asp-Glu-Glu Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O GHODABZPVZMWCE-FXQIFTODSA-N 0.000 description 6
- LFQSCWFLJHTTHZ-UHFFFAOYSA-N Ethanol Chemical compound CCO LFQSCWFLJHTTHZ-UHFFFAOYSA-N 0.000 description 6
- ATVYZJGOZLVXDK-IUCAKERBSA-N Glu-Leu-Gly Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O ATVYZJGOZLVXDK-IUCAKERBSA-N 0.000 description 6
- WHUUTDBJXJRKMK-UHFFFAOYSA-N Glutamic acid Natural products OC(=O)C(N)CCC(O)=O WHUUTDBJXJRKMK-UHFFFAOYSA-N 0.000 description 6
- 108010065920 Insulin Lispro Proteins 0.000 description 6
- 108010002311 N-glycylglutamic acid Proteins 0.000 description 6
- 108700019146 Transgenes Proteins 0.000 description 6
- 108010055341 glutamyl-glutamic acid Proteins 0.000 description 6
- 229930027917 kanamycin Natural products 0.000 description 6
- 229960000318 kanamycin Drugs 0.000 description 6
- SBUJHOSQTJFQJX-NOAMYHISSA-N kanamycin Chemical compound O[C@@H]1[C@@H](O)[C@H](O)[C@@H](CN)O[C@@H]1O[C@H]1[C@H](O)[C@@H](O[C@@H]2[C@@H]([C@@H](N)[C@H](O)[C@@H](CO)O2)O)[C@H](N)C[C@@H]1N SBUJHOSQTJFQJX-NOAMYHISSA-N 0.000 description 6
- 229930182823 kanamycin A Natural products 0.000 description 6
- 239000002609 medium Substances 0.000 description 6
- 108091032973 (ribonucleotides)n+m Proteins 0.000 description 5
- 241000589155 Agrobacterium tumefaciens Species 0.000 description 5
- NNQDRRUXFJYCCJ-NHCYSSNCSA-N Glu-Pro-Val Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(O)=O NNQDRRUXFJYCCJ-NHCYSSNCSA-N 0.000 description 5
- IMAKMJCBYCSMHM-AVGNSLFASA-N Lys-Glu-Lys Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CCCCN IMAKMJCBYCSMHM-AVGNSLFASA-N 0.000 description 5
- SBVPYBFMIGDIDX-SRVKXCTJSA-N Pro-Pro-Pro Chemical compound OC(=O)[C@@H]1CCCN1C(=O)[C@H]1N(C(=O)[C@H]2NCCC2)CCC1 SBVPYBFMIGDIDX-SRVKXCTJSA-N 0.000 description 5
- 230000002159 abnormal effect Effects 0.000 description 5
- 108010047495 alanylglycine Proteins 0.000 description 5
- 108010013835 arginine glutamate Proteins 0.000 description 5
- 238000006243 chemical reaction Methods 0.000 description 5
- 238000010276 construction Methods 0.000 description 5
- 239000012634 fragment Substances 0.000 description 5
- 238000003752 polymerase chain reaction Methods 0.000 description 5
- 230000008569 process Effects 0.000 description 5
- 230000000644 propagated effect Effects 0.000 description 5
- UUTKICFRNVKFRG-WDSKDSINSA-N (4R)-3-[oxo-[(2S)-5-oxo-2-pyrrolidinyl]methyl]-4-thiazolidinecarboxylic acid Chemical compound OC(=O)[C@@H]1CSCN1C(=O)[C@H]1NC(=O)CC1 UUTKICFRNVKFRG-WDSKDSINSA-N 0.000 description 4
- JBGSZRYCXBPWGX-BQBZGAKWSA-N Ala-Arg-Gly Chemical compound OC(=O)CNC(=O)[C@@H](NC(=O)[C@@H](N)C)CCCN=C(N)N JBGSZRYCXBPWGX-BQBZGAKWSA-N 0.000 description 4
- HUUOZYZWNCXTFK-INTQDDNPSA-N Ala-His-Pro Chemical compound C[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N2CCC[C@@H]2C(=O)O)N HUUOZYZWNCXTFK-INTQDDNPSA-N 0.000 description 4
- WEZNQZHACPSMEF-QEJZJMRPSA-N Ala-Phe-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)C)CC1=CC=CC=C1 WEZNQZHACPSMEF-QEJZJMRPSA-N 0.000 description 4
- IYKVSFNGSWTTNZ-GUBZILKMSA-N Ala-Val-Arg Chemical compound C[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CCCN=C(N)N IYKVSFNGSWTTNZ-GUBZILKMSA-N 0.000 description 4
- 108700040922 Arabidopsis LFY Proteins 0.000 description 4
- JGDGLDNAQJJGJI-AVGNSLFASA-N Arg-Arg-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@H](CCCN=C(N)N)N JGDGLDNAQJJGJI-AVGNSLFASA-N 0.000 description 4
- QAXCZGMLVICQKS-SRVKXCTJSA-N Arg-Glu-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)O)NC(=O)[C@H](CCCN=C(N)N)N QAXCZGMLVICQKS-SRVKXCTJSA-N 0.000 description 4
- AUFHLLPVPSMEOG-YUMQZZPRSA-N Arg-Gly-Glu Chemical compound NC(N)=NCCC[C@H](N)C(=O)NCC(=O)N[C@@H](CCC(O)=O)C(O)=O AUFHLLPVPSMEOG-YUMQZZPRSA-N 0.000 description 4
- COXMUHNBYCVVRG-DCAQKATOSA-N Arg-Leu-Ser Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O COXMUHNBYCVVRG-DCAQKATOSA-N 0.000 description 4
- NVPHRWNWTKYIST-BPNCWPANSA-N Arg-Tyr-Ala Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](C)C(O)=O)CC1=CC=C(O)C=C1 NVPHRWNWTKYIST-BPNCWPANSA-N 0.000 description 4
- JWKDQOORUCYUIW-ZPFDUUQYSA-N Asn-Lys-Ile Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O JWKDQOORUCYUIW-ZPFDUUQYSA-N 0.000 description 4
- ZAESWDKAMDVHLL-RCOVLWMOSA-N Asn-Val-Gly Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](C(C)C)C(=O)NCC(O)=O ZAESWDKAMDVHLL-RCOVLWMOSA-N 0.000 description 4
- TZOZNVLBTAFJRW-UGYAYLCHSA-N Asp-Ile-Asp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)[C@H](CC(=O)O)N TZOZNVLBTAFJRW-UGYAYLCHSA-N 0.000 description 4
- 240000007124 Brassica oleracea Species 0.000 description 4
- 235000011299 Brassica oleracea var botrytis Nutrition 0.000 description 4
- 240000003259 Brassica oleracea var. botrytis Species 0.000 description 4
- BLGNLNRBABWDST-CIUDSAMLSA-N Cys-Leu-Asp Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)[C@H](CS)N BLGNLNRBABWDST-CIUDSAMLSA-N 0.000 description 4
- TXGDWPBLUFQODU-XGEHTFHBSA-N Cys-Pro-Thr Chemical compound [H]N[C@@H](CS)C(=O)N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)O)C(O)=O TXGDWPBLUFQODU-XGEHTFHBSA-N 0.000 description 4
- DTCCMDYODDPHBG-ACZMJKKPSA-N Gln-Ala-Cys Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C)C(=O)N[C@@H](CS)C(O)=O DTCCMDYODDPHBG-ACZMJKKPSA-N 0.000 description 4
- KWUSGAIFNHQCBY-DCAQKATOSA-N Gln-Arg-Arg Chemical compound NC(=O)CC[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O KWUSGAIFNHQCBY-DCAQKATOSA-N 0.000 description 4
- PCKOTDPDHIBGRW-CIUDSAMLSA-N Gln-Cys-Arg Chemical compound C(C[C@@H](C(=O)O)NC(=O)[C@H](CS)NC(=O)[C@H](CCC(=O)N)N)CN=C(N)N PCKOTDPDHIBGRW-CIUDSAMLSA-N 0.000 description 4
- PKVWNYGXMNWJSI-CIUDSAMLSA-N Gln-Gln-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O PKVWNYGXMNWJSI-CIUDSAMLSA-N 0.000 description 4
- IULKWYSYZSURJK-AVGNSLFASA-N Gln-Leu-Lys Chemical compound NC(=O)CC[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(O)=O IULKWYSYZSURJK-AVGNSLFASA-N 0.000 description 4
- ZFBBMCKQSNJZSN-AUTRQRHGSA-N Gln-Val-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O ZFBBMCKQSNJZSN-AUTRQRHGSA-N 0.000 description 4
- ZMXZGYLINVNTKH-DZKIICNBSA-N Gln-Val-Phe Chemical compound NC(=O)CC[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 ZMXZGYLINVNTKH-DZKIICNBSA-N 0.000 description 4
- BUZMZDDKFCSKOT-CIUDSAMLSA-N Glu-Glu-Glu Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O BUZMZDDKFCSKOT-CIUDSAMLSA-N 0.000 description 4
- ZQIMMEYPEXIYBB-IUCAKERBSA-N Gly-Glu-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CCC(O)=O)NC(=O)CN ZQIMMEYPEXIYBB-IUCAKERBSA-N 0.000 description 4
- JSNNHGHYGYMVCK-XVKPBYJWSA-N Gly-Glu-Val Chemical compound [H]NCC(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O JSNNHGHYGYMVCK-XVKPBYJWSA-N 0.000 description 4
- ITZOBNKQDZEOCE-NHCYSSNCSA-N Gly-Ile-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)CN ITZOBNKQDZEOCE-NHCYSSNCSA-N 0.000 description 4
- 206010020649 Hyperkeratosis Diseases 0.000 description 4
- FJWYJQRCVNGEAQ-ZPFDUUQYSA-N Ile-Asn-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CCCCN)C(=O)O)N FJWYJQRCVNGEAQ-ZPFDUUQYSA-N 0.000 description 4
- MITYXXNZSZLHGG-OBAATPRFSA-N Ile-Trp-Tyr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)N[C@@H](CC3=CC=C(C=C3)O)C(=O)O)N MITYXXNZSZLHGG-OBAATPRFSA-N 0.000 description 4
- NTRAGDHVSGKUSF-AVGNSLFASA-N Leu-Arg-Arg Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O NTRAGDHVSGKUSF-AVGNSLFASA-N 0.000 description 4
- QLQHWWCSCLZUMA-KKUMJFAQSA-N Leu-Asp-Tyr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 QLQHWWCSCLZUMA-KKUMJFAQSA-N 0.000 description 4
- LJKJVTCIRDCITR-SRVKXCTJSA-N Leu-Cys-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N LJKJVTCIRDCITR-SRVKXCTJSA-N 0.000 description 4
- WIDZHJTYKYBLSR-DCAQKATOSA-N Leu-Glu-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O WIDZHJTYKYBLSR-DCAQKATOSA-N 0.000 description 4
- KTOIECMYZZGVSI-BZSNNMDCSA-N Leu-Phe-His Chemical compound C([C@H](NC(=O)[C@@H](N)CC(C)C)C(=O)N[C@@H](CC=1NC=NC=1)C(O)=O)C1=CC=CC=C1 KTOIECMYZZGVSI-BZSNNMDCSA-N 0.000 description 4
- VJGQRELPQWNURN-JYJNAYRXSA-N Leu-Tyr-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(O)=O)C(O)=O VJGQRELPQWNURN-JYJNAYRXSA-N 0.000 description 4
- AAKRWBIIGKPOKQ-ONGXEEELSA-N Leu-Val-Gly Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)NCC(O)=O AAKRWBIIGKPOKQ-ONGXEEELSA-N 0.000 description 4
- SLQJJFAVWSZLBL-BJDJZHNGSA-N Lys-Ile-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H]([C@@H](C)CC)NC(=O)[C@@H](N)CCCCN SLQJJFAVWSZLBL-BJDJZHNGSA-N 0.000 description 4
- OVAOHZIOUBEQCJ-IHRRRGAJSA-N Lys-Leu-Arg Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O OVAOHZIOUBEQCJ-IHRRRGAJSA-N 0.000 description 4
- RIJCHEVHFWMDKD-SRVKXCTJSA-N Lys-Lys-Asn Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(N)=O)C(O)=O RIJCHEVHFWMDKD-SRVKXCTJSA-N 0.000 description 4
- DNDVVILEHVMWIS-LPEHRKFASA-N Met-Asp-Pro Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N DNDVVILEHVMWIS-LPEHRKFASA-N 0.000 description 4
- FYRUJIJAUPHUNB-IUCAKERBSA-N Met-Gly-Arg Chemical compound CSCC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCNC(N)=N FYRUJIJAUPHUNB-IUCAKERBSA-N 0.000 description 4
- VAGCEUUEMMXFEX-GUBZILKMSA-N Met-Met-Asn Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(N)=O)C(O)=O VAGCEUUEMMXFEX-GUBZILKMSA-N 0.000 description 4
- RORUIHAWOLADSH-HJWJTTGWSA-N Phe-Ile-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H]([C@@H](C)CC)NC(=O)[C@@H](N)CC1=CC=CC=C1 RORUIHAWOLADSH-HJWJTTGWSA-N 0.000 description 4
- PUQRDHNIOONJJN-AVGNSLFASA-N Pro-Lys-Met Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCSC)C(O)=O PUQRDHNIOONJJN-AVGNSLFASA-N 0.000 description 4
- 108700005075 Regulator Genes Proteins 0.000 description 4
- XVAUJOAYHWWNQF-ZLUOBGJFSA-N Ser-Asn-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](C)C(O)=O XVAUJOAYHWWNQF-ZLUOBGJFSA-N 0.000 description 4
- KBLYJPQSNGTDIU-LOKLDPHHSA-N Thr-Glu-Pro Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N)O KBLYJPQSNGTDIU-LOKLDPHHSA-N 0.000 description 4
- XOWKUMFHEZLKLT-CIQUZCHMSA-N Thr-Ile-Ala Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C)C(O)=O XOWKUMFHEZLKLT-CIQUZCHMSA-N 0.000 description 4
- 108091023040 Transcription factor Proteins 0.000 description 4
- 102000040945 Transcription factor Human genes 0.000 description 4
- TVOGEPLDNYTAHD-CQDKDKBSSA-N Tyr-Ala-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@@H](N)CC1=CC=C(O)C=C1 TVOGEPLDNYTAHD-CQDKDKBSSA-N 0.000 description 4
- KGSDLCMCDFETHU-YESZJQIVSA-N Tyr-Lys-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCCCN)NC(=O)[C@H](CC2=CC=C(C=C2)O)N)C(=O)O KGSDLCMCDFETHU-YESZJQIVSA-N 0.000 description 4
- XUIOBCQESNDTDE-FQPOAREZSA-N Tyr-Thr-Ala Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](C)C(=O)O)NC(=O)[C@H](CC1=CC=C(C=C1)O)N)O XUIOBCQESNDTDE-FQPOAREZSA-N 0.000 description 4
- PIFJAFRUVWZRKR-QMMMGPOBSA-N Val-Gly-Gly Chemical compound CC(C)[C@H]([NH3+])C(=O)NCC(=O)NCC([O-])=O PIFJAFRUVWZRKR-QMMMGPOBSA-N 0.000 description 4
- MANXHLOVEUHVFD-DCAQKATOSA-N Val-His-Cys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CS)C(=O)O)N MANXHLOVEUHVFD-DCAQKATOSA-N 0.000 description 4
- MIKHIIQMRFYVOR-RCWTZXSCSA-N Val-Pro-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@@H]1CCCN1C(=O)[C@H](C(C)C)N)O MIKHIIQMRFYVOR-RCWTZXSCSA-N 0.000 description 4
- PQSNETRGCRUOGP-KKHAAJSZSA-N Val-Thr-Asn Chemical compound CC(C)[C@H](N)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@H](C(O)=O)CC(N)=O PQSNETRGCRUOGP-KKHAAJSZSA-N 0.000 description 4
- 235000002017 Zea mays subsp mays Nutrition 0.000 description 4
- 241000482268 Zea mays subsp. mays Species 0.000 description 4
- KOSRFJWDECSPRO-UHFFFAOYSA-N alpha-L-glutamyl-L-glutamic acid Natural products OC(=O)CCC(N)C(=O)NC(CCC(O)=O)C(O)=O KOSRFJWDECSPRO-UHFFFAOYSA-N 0.000 description 4
- 108010018691 arginyl-threonyl-arginine Proteins 0.000 description 4
- 108010040443 aspartyl-aspartic acid Proteins 0.000 description 4
- 108010038633 aspartylglutamate Proteins 0.000 description 4
- 238000009395 breeding Methods 0.000 description 4
- 230000001488 breeding effect Effects 0.000 description 4
- 230000008859 change Effects 0.000 description 4
- 230000001276 controlling effect Effects 0.000 description 4
- 230000002068 genetic effect Effects 0.000 description 4
- 108010057083 glutamyl-aspartyl-leucine Proteins 0.000 description 4
- VPZXBVLAVMBEQI-UHFFFAOYSA-N glycyl-DL-alpha-alanine Natural products OC(=O)C(C)NC(=O)CN VPZXBVLAVMBEQI-UHFFFAOYSA-N 0.000 description 4
- 108010067216 glycyl-glycyl-glycine Proteins 0.000 description 4
- 108010066198 glycyl-leucyl-phenylalanine Proteins 0.000 description 4
- 108010015792 glycyllysine Proteins 0.000 description 4
- 108010085325 histidylproline Proteins 0.000 description 4
- 230000000977 initiatory effect Effects 0.000 description 4
- 108010051673 leucyl-glycyl-phenylalanine Proteins 0.000 description 4
- 108010025153 lysyl-alanyl-alanine Proteins 0.000 description 4
- 108010018625 phenylalanylarginine Proteins 0.000 description 4
- 108010004914 prolylarginine Proteins 0.000 description 4
- 108010029020 prolylglycine Proteins 0.000 description 4
- 239000000523 sample Substances 0.000 description 4
- 238000012216 screening Methods 0.000 description 4
- 108010071207 serylmethionine Proteins 0.000 description 4
- 239000000725 suspension Substances 0.000 description 4
- 238000012546 transfer Methods 0.000 description 4
- QTBSBXVTEAMEQO-UHFFFAOYSA-N Acetic acid Chemical compound CC(O)=O QTBSBXVTEAMEQO-UHFFFAOYSA-N 0.000 description 3
- AWZKCUCQJNTBAD-SRVKXCTJSA-N Ala-Leu-Lys Chemical compound C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CCCCN AWZKCUCQJNTBAD-SRVKXCTJSA-N 0.000 description 3
- 241000219195 Arabidopsis thaliana Species 0.000 description 3
- LMPKCSXZJSXBBL-NHCYSSNCSA-N Arg-Gln-Val Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(O)=O LMPKCSXZJSXBBL-NHCYSSNCSA-N 0.000 description 3
- PLOKOIJSGCISHE-BYULHYEWSA-N Asp-Val-Asn Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(N)=O)C(O)=O PLOKOIJSGCISHE-BYULHYEWSA-N 0.000 description 3
- 235000003899 Brassica oleracea var acephala Nutrition 0.000 description 3
- 241000219193 Brassicaceae Species 0.000 description 3
- 108020004705 Codon Proteins 0.000 description 3
- VNLYIYOYUNGURO-ZLUOBGJFSA-N Cys-Asp-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H](CS)N VNLYIYOYUNGURO-ZLUOBGJFSA-N 0.000 description 3
- WSFSSNUMVMOOMR-UHFFFAOYSA-N Formaldehyde Chemical compound O=C WSFSSNUMVMOOMR-UHFFFAOYSA-N 0.000 description 3
- CGVWDTRDPLOMHZ-FXQIFTODSA-N Gln-Glu-Asp Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O CGVWDTRDPLOMHZ-FXQIFTODSA-N 0.000 description 3
- XHWLNISLUFEWNS-CIUDSAMLSA-N Glu-Gln-Gln Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O XHWLNISLUFEWNS-CIUDSAMLSA-N 0.000 description 3
- KIEICAOUSNYOLM-NRPADANISA-N Glu-Val-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](C)C(O)=O KIEICAOUSNYOLM-NRPADANISA-N 0.000 description 3
- MHXKHKWHPNETGG-QWRGUYRKSA-N Gly-Lys-Leu Chemical compound [H]NCC(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(O)=O MHXKHKWHPNETGG-QWRGUYRKSA-N 0.000 description 3
- 108700005087 Homeobox Genes Proteins 0.000 description 3
- HOLOYAZCIHDQNS-YVNDNENWSA-N Ile-Gln-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N HOLOYAZCIHDQNS-YVNDNENWSA-N 0.000 description 3
- CLVUXCBGKUECIT-HJGDQZAQSA-N Leu-Asp-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O CLVUXCBGKUECIT-HJGDQZAQSA-N 0.000 description 3
- RVVBWTWPNFDYBE-SRVKXCTJSA-N Leu-Glu-Arg Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O RVVBWTWPNFDYBE-SRVKXCTJSA-N 0.000 description 3
- KPYAOIVPJKPIOU-KKUMJFAQSA-N Leu-Lys-Lys Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(O)=O KPYAOIVPJKPIOU-KKUMJFAQSA-N 0.000 description 3
- CLBGMWIYPYAZPR-AVGNSLFASA-N Lys-Arg-Arg Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O CLBGMWIYPYAZPR-AVGNSLFASA-N 0.000 description 3
- DRCILAJNUJKAHC-SRVKXCTJSA-N Lys-Glu-Arg Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O DRCILAJNUJKAHC-SRVKXCTJSA-N 0.000 description 3
- AUEJLPRZGVVDNU-UHFFFAOYSA-N N-L-tyrosyl-L-leucine Natural products CC(C)CC(C(O)=O)NC(=O)C(N)CC1=CC=C(O)C=C1 AUEJLPRZGVVDNU-UHFFFAOYSA-N 0.000 description 3
- 108700026244 Open Reading Frames Proteins 0.000 description 3
- UAMFZRNCIFFMLE-FHWLQOOXSA-N Phe-Glu-Tyr Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CC2=CC=C(C=C2)O)C(=O)O)N UAMFZRNCIFFMLE-FHWLQOOXSA-N 0.000 description 3
- UCOYFSCEIWQYNL-FXQIFTODSA-N Ser-Cys-Met Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CS)C(=O)N[C@@H](CCSC)C(O)=O UCOYFSCEIWQYNL-FXQIFTODSA-N 0.000 description 3
- WUXCHQZLUHBSDJ-LKXGYXEUSA-N Ser-Thr-Asp Chemical compound OC[C@H](N)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@@H](CC(O)=O)C(O)=O WUXCHQZLUHBSDJ-LKXGYXEUSA-N 0.000 description 3
- YEDSOSIKVUMIJE-DCAQKATOSA-N Ser-Val-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O YEDSOSIKVUMIJE-DCAQKATOSA-N 0.000 description 3
- 241000208292 Solanaceae Species 0.000 description 3
- NAXBBCLCEOTAIG-RHYQMDGZSA-N Thr-Arg-Lys Chemical compound NC(N)=NCCC[C@H](NC(=O)[C@@H](N)[C@H](O)C)C(=O)N[C@@H](CCCCN)C(O)=O NAXBBCLCEOTAIG-RHYQMDGZSA-N 0.000 description 3
- ABWNZPOIUJMNKT-IXOXFDKPSA-N Thr-Phe-Ser Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CO)C(O)=O ABWNZPOIUJMNKT-IXOXFDKPSA-N 0.000 description 3
- 150000001413 amino acids Chemical class 0.000 description 3
- 238000004458 analytical method Methods 0.000 description 3
- 229910002056 binary alloy Inorganic materials 0.000 description 3
- 230000003115 biocidal effect Effects 0.000 description 3
- 230000000295 complement effect Effects 0.000 description 3
- 235000013399 edible fruits Nutrition 0.000 description 3
- 238000005516 engineering process Methods 0.000 description 3
- 230000007613 environmental effect Effects 0.000 description 3
- 238000012239 gene modification Methods 0.000 description 3
- 230000005017 genetic modification Effects 0.000 description 3
- 235000013617 genetically modified food Nutrition 0.000 description 3
- 239000000411 inducer Substances 0.000 description 3
- 108010064235 lysylglycine Proteins 0.000 description 3
- 108020004999 messenger RNA Proteins 0.000 description 3
- 108010058731 nopaline synthase Proteins 0.000 description 3
- 230000001172 regenerating effect Effects 0.000 description 3
- 238000012552 review Methods 0.000 description 3
- 108010048818 seryl-histidine Proteins 0.000 description 3
- 238000003786 synthesis reaction Methods 0.000 description 3
- OZRFYUJEXYKQDV-UHFFFAOYSA-N 2-[[2-[[2-[(2-amino-3-carboxypropanoyl)amino]-3-carboxypropanoyl]amino]-3-carboxypropanoyl]amino]butanedioic acid Chemical compound OC(=O)CC(N)C(=O)NC(CC(O)=O)C(=O)NC(CC(O)=O)C(=O)NC(CC(O)=O)C(O)=O OZRFYUJEXYKQDV-UHFFFAOYSA-N 0.000 description 2
- HHGYNJRJIINWAK-FXQIFTODSA-N Ala-Ala-Arg Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCCN=C(N)N HHGYNJRJIINWAK-FXQIFTODSA-N 0.000 description 2
- RLMISHABBKUNFO-WHFBIAKZSA-N Ala-Ala-Gly Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)NCC(O)=O RLMISHABBKUNFO-WHFBIAKZSA-N 0.000 description 2
- JBVSSSZFNTXJDX-YTLHQDLWSA-N Ala-Ala-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@H](C)N JBVSSSZFNTXJDX-YTLHQDLWSA-N 0.000 description 2
- LZRNYBIJOSKKRJ-XVYDVKMFSA-N Ala-Asp-His Chemical compound C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N LZRNYBIJOSKKRJ-XVYDVKMFSA-N 0.000 description 2
- WJRXVTCKASUIFF-FXQIFTODSA-N Ala-Cys-Arg Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CS)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O WJRXVTCKASUIFF-FXQIFTODSA-N 0.000 description 2
- RXTBLQVXNIECFP-FXQIFTODSA-N Ala-Gln-Gln Chemical compound C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O RXTBLQVXNIECFP-FXQIFTODSA-N 0.000 description 2
- KXEVYGKATAMXJJ-ACZMJKKPSA-N Ala-Glu-Asp Chemical compound C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O KXEVYGKATAMXJJ-ACZMJKKPSA-N 0.000 description 2
- VGPWRRFOPXVGOH-BYPYZUCNSA-N Ala-Gly-Gly Chemical compound C[C@H](N)C(=O)NCC(=O)NCC(O)=O VGPWRRFOPXVGOH-BYPYZUCNSA-N 0.000 description 2
- LBYMZCVBOKYZNS-CIUDSAMLSA-N Ala-Leu-Asp Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O LBYMZCVBOKYZNS-CIUDSAMLSA-N 0.000 description 2
- SOBIAADAMRHGKH-CIUDSAMLSA-N Ala-Leu-Ser Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O SOBIAADAMRHGKH-CIUDSAMLSA-N 0.000 description 2
- OLVCTPPSXNRGKV-GUBZILKMSA-N Ala-Pro-Pro Chemical compound C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N1[C@H](C(O)=O)CCC1 OLVCTPPSXNRGKV-GUBZILKMSA-N 0.000 description 2
- XWFWAXPOLRTDFZ-FXQIFTODSA-N Ala-Pro-Ser Chemical compound C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O XWFWAXPOLRTDFZ-FXQIFTODSA-N 0.000 description 2
- OMCKWYSDUQBYCN-FXQIFTODSA-N Ala-Ser-Met Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCSC)C(O)=O OMCKWYSDUQBYCN-FXQIFTODSA-N 0.000 description 2
- AETQNIIFKCMVHP-UVBJJODRSA-N Ala-Trp-Arg Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O AETQNIIFKCMVHP-UVBJJODRSA-N 0.000 description 2
- PGNNQOJOEGFAOR-KWQFWETISA-N Ala-Tyr-Gly Chemical compound OC(=O)CNC(=O)[C@@H](NC(=O)[C@@H](N)C)CC1=CC=C(O)C=C1 PGNNQOJOEGFAOR-KWQFWETISA-N 0.000 description 2
- 241000207875 Antirrhinum Species 0.000 description 2
- GXCSUJQOECMKPV-CIUDSAMLSA-N Arg-Ala-Gln Chemical compound C[C@H](NC(=O)[C@@H](N)CCCNC(N)=N)C(=O)N[C@@H](CCC(N)=O)C(O)=O GXCSUJQOECMKPV-CIUDSAMLSA-N 0.000 description 2
- BHSYMWWMVRPCPA-CYDGBPFRSA-N Arg-Arg-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@@H](N)CCCN=C(N)N BHSYMWWMVRPCPA-CYDGBPFRSA-N 0.000 description 2
- RVDVDRUZWZIBJQ-CIUDSAMLSA-N Arg-Asn-Glu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O RVDVDRUZWZIBJQ-CIUDSAMLSA-N 0.000 description 2
- NUBPTCMEOCKWDO-DCAQKATOSA-N Arg-Asn-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](CCCN=C(N)N)N NUBPTCMEOCKWDO-DCAQKATOSA-N 0.000 description 2
- MMGCRPZQZWTZTA-IHRRRGAJSA-N Arg-His-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N MMGCRPZQZWTZTA-IHRRRGAJSA-N 0.000 description 2
- OOIMKQRCPJBGPD-XUXIUFHCSA-N Arg-Ile-Leu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(C)C)C(O)=O OOIMKQRCPJBGPD-XUXIUFHCSA-N 0.000 description 2
- GMFAGHNRXPSSJS-SRVKXCTJSA-N Arg-Leu-Gln Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O GMFAGHNRXPSSJS-SRVKXCTJSA-N 0.000 description 2
- YBZMTKUDWXZLIX-UWVGGRQHSA-N Arg-Leu-Gly Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O YBZMTKUDWXZLIX-UWVGGRQHSA-N 0.000 description 2
- SSZGOKWBHLOCHK-DCAQKATOSA-N Arg-Lys-Asn Chemical compound NC(=O)C[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CCCN=C(N)N SSZGOKWBHLOCHK-DCAQKATOSA-N 0.000 description 2
- BTJVOUQWFXABOI-IHRRRGAJSA-N Arg-Lys-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CCCNC(N)=N BTJVOUQWFXABOI-IHRRRGAJSA-N 0.000 description 2
- WKPXXXUSUHAXDE-SRVKXCTJSA-N Arg-Pro-Arg Chemical compound NC(N)=NCCC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCCN=C(N)N)C(O)=O WKPXXXUSUHAXDE-SRVKXCTJSA-N 0.000 description 2
- VENMDXUVHSKEIN-GUBZILKMSA-N Arg-Ser-Arg Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O VENMDXUVHSKEIN-GUBZILKMSA-N 0.000 description 2
- ASQKVGRCKOFKIU-KZVJFYERSA-N Arg-Thr-Ala Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](C)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N)O ASQKVGRCKOFKIU-KZVJFYERSA-N 0.000 description 2
- XWGJDUSDTRPQRK-ZLUOBGJFSA-N Asn-Ala-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@@H](N)CC(N)=O XWGJDUSDTRPQRK-ZLUOBGJFSA-N 0.000 description 2
- MFFOYNGMOYFPBD-DCAQKATOSA-N Asn-Arg-Leu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(C)C)C(O)=O MFFOYNGMOYFPBD-DCAQKATOSA-N 0.000 description 2
- ZZXMOQIUIJJOKZ-ZLUOBGJFSA-N Asn-Asn-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CC(N)=O)NC(=O)[C@@H](N)CC(N)=O ZZXMOQIUIJJOKZ-ZLUOBGJFSA-N 0.000 description 2
- OLISTMZJGQUOGS-GMOBBJLQSA-N Asn-Ile-Arg Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)NC(=O)[C@H](CC(=O)N)N OLISTMZJGQUOGS-GMOBBJLQSA-N 0.000 description 2
- JBDLMLZNDRLDIX-HJGDQZAQSA-N Asn-Thr-Leu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(O)=O JBDLMLZNDRLDIX-HJGDQZAQSA-N 0.000 description 2
- ULZOQOKFYMXHPZ-AQZXSJQPSA-N Asn-Trp-Thr Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H]([C@@H](C)O)C(O)=O ULZOQOKFYMXHPZ-AQZXSJQPSA-N 0.000 description 2
- KRXIWXCXOARFNT-ZLUOBGJFSA-N Asp-Ala-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](C)NC(=O)[C@@H](N)CC(O)=O KRXIWXCXOARFNT-ZLUOBGJFSA-N 0.000 description 2
- PBVLJOIPOGUQQP-CIUDSAMLSA-N Asp-Ala-Leu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(O)=O PBVLJOIPOGUQQP-CIUDSAMLSA-N 0.000 description 2
- NECWUSYTYSIFNC-DLOVCJGASA-N Asp-Ala-Phe Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 NECWUSYTYSIFNC-DLOVCJGASA-N 0.000 description 2
- VPSHHQXIWLGVDD-ZLUOBGJFSA-N Asp-Asp-Asp Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O VPSHHQXIWLGVDD-ZLUOBGJFSA-N 0.000 description 2
- WCFCYFDBMNFSPA-ACZMJKKPSA-N Asp-Asp-Glu Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CCC(O)=O WCFCYFDBMNFSPA-ACZMJKKPSA-N 0.000 description 2
- SBHUBSDEZQFJHJ-CIUDSAMLSA-N Asp-Asp-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(O)=O)NC(=O)[C@@H](N)CC(O)=O SBHUBSDEZQFJHJ-CIUDSAMLSA-N 0.000 description 2
- KIJLEFNHWSXHRU-NUMRIWBASA-N Asp-Gln-Thr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O KIJLEFNHWSXHRU-NUMRIWBASA-N 0.000 description 2
- XJQRWGXKUSDEFI-ACZMJKKPSA-N Asp-Glu-Asn Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O XJQRWGXKUSDEFI-ACZMJKKPSA-N 0.000 description 2
- PAYPSKIBMDHZPI-CIUDSAMLSA-N Asp-Leu-Asp Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O PAYPSKIBMDHZPI-CIUDSAMLSA-N 0.000 description 2
- LTCKTLYKRMCFOC-KKUMJFAQSA-N Asp-Phe-Leu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(C)C)C(O)=O LTCKTLYKRMCFOC-KKUMJFAQSA-N 0.000 description 2
- KGHLGJAXYSVNJP-WHFBIAKZSA-N Asp-Ser-Gly Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CO)C(=O)NCC(O)=O KGHLGJAXYSVNJP-WHFBIAKZSA-N 0.000 description 2
- RSMZEHCMIOKNMW-GSSVUCPTSA-N Asp-Thr-Thr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O RSMZEHCMIOKNMW-GSSVUCPTSA-N 0.000 description 2
- LEYKQPDPZJIRTA-AQZXSJQPSA-N Asp-Trp-Thr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H]([C@@H](C)O)C(O)=O LEYKQPDPZJIRTA-AQZXSJQPSA-N 0.000 description 2
- 240000002791 Brassica napus Species 0.000 description 2
- 235000011301 Brassica oleracea var capitata Nutrition 0.000 description 2
- 235000017647 Brassica oleracea var italica Nutrition 0.000 description 2
- 235000001169 Brassica oleracea var oleracea Nutrition 0.000 description 2
- 101100382321 Caenorhabditis elegans cal-1 gene Proteins 0.000 description 2
- 240000004160 Capsicum annuum Species 0.000 description 2
- 235000008534 Capsicum annuum var annuum Nutrition 0.000 description 2
- 244000241257 Cucumis melo Species 0.000 description 2
- DZLQXIFVQFTFJY-BYPYZUCNSA-N Cys-Gly-Gly Chemical compound SC[C@H](N)C(=O)NCC(=O)NCC(O)=O DZLQXIFVQFTFJY-BYPYZUCNSA-N 0.000 description 2
- RRJOQIBQVZDVCW-SRVKXCTJSA-N Cys-His-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CC1=CN=CN1)NC(=O)[C@H](CS)N RRJOQIBQVZDVCW-SRVKXCTJSA-N 0.000 description 2
- 108010066133 D-octopine dehydrogenase Proteins 0.000 description 2
- 230000004568 DNA-binding Effects 0.000 description 2
- TWHDOEYLXXQYOZ-FXQIFTODSA-N Gln-Asn-Gln Chemical compound C(CC(=O)N)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N TWHDOEYLXXQYOZ-FXQIFTODSA-N 0.000 description 2
- GMGKDVVBSVVKCT-NUMRIWBASA-N Gln-Asn-Thr Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O GMGKDVVBSVVKCT-NUMRIWBASA-N 0.000 description 2
- XFKUFUJECJUQTQ-CIUDSAMLSA-N Gln-Gln-Glu Chemical compound NC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O XFKUFUJECJUQTQ-CIUDSAMLSA-N 0.000 description 2
- ZQPOVSJFBBETHQ-CIUDSAMLSA-N Gln-Glu-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O ZQPOVSJFBBETHQ-CIUDSAMLSA-N 0.000 description 2
- NXPXQIZKDOXIHH-JSGCOSHPSA-N Gln-Gly-Trp Chemical compound C1=CC=C2C(=C1)C(=CN2)C[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CCC(=O)N)N NXPXQIZKDOXIHH-JSGCOSHPSA-N 0.000 description 2
- MWERYIXRDZDXOA-QEWYBTABSA-N Gln-Ile-Phe Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O MWERYIXRDZDXOA-QEWYBTABSA-N 0.000 description 2
- XFAUJGNLHIGXET-AVGNSLFASA-N Gln-Leu-Leu Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O XFAUJGNLHIGXET-AVGNSLFASA-N 0.000 description 2
- YPFFHGRJCUBXPX-NHCYSSNCSA-N Gln-Pro-Val Chemical compound CC(C)[C@H](NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)CCC(N)=O)C(O)=O YPFFHGRJCUBXPX-NHCYSSNCSA-N 0.000 description 2
- UBRQJXFDVZNYJP-AVGNSLFASA-N Gln-Tyr-Ser Chemical compound C1=CC(=CC=C1C[C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](CCC(=O)N)N)O UBRQJXFDVZNYJP-AVGNSLFASA-N 0.000 description 2
- RUFHOVYUYSNDNY-ACZMJKKPSA-N Glu-Ala-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](C)NC(=O)[C@@H](N)CCC(O)=O RUFHOVYUYSNDNY-ACZMJKKPSA-N 0.000 description 2
- QPRZKNOOOBWXSU-CIUDSAMLSA-N Glu-Asp-Arg Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CCCN=C(N)N QPRZKNOOOBWXSU-CIUDSAMLSA-N 0.000 description 2
- CGOHAEBMDSEKFB-FXQIFTODSA-N Glu-Glu-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O CGOHAEBMDSEKFB-FXQIFTODSA-N 0.000 description 2
- NKLRYVLERDYDBI-FXQIFTODSA-N Glu-Glu-Asp Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O NKLRYVLERDYDBI-FXQIFTODSA-N 0.000 description 2
- AIGROOHQXCACHL-WDSKDSINSA-N Glu-Gly-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)NCC(=O)N[C@@H](C)C(O)=O AIGROOHQXCACHL-WDSKDSINSA-N 0.000 description 2
- OGNJZUXUTPQVBR-BQBZGAKWSA-N Glu-Gly-Glu Chemical compound OC(=O)CC[C@H](N)C(=O)NCC(=O)N[C@@H](CCC(O)=O)C(O)=O OGNJZUXUTPQVBR-BQBZGAKWSA-N 0.000 description 2
- ZWQVYZXPYSYPJD-RYUDHWBXSA-N Glu-Gly-Phe Chemical compound OC(=O)CC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 ZWQVYZXPYSYPJD-RYUDHWBXSA-N 0.000 description 2
- XOFYVODYSNKPDK-AVGNSLFASA-N Glu-His-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)NC(=O)[C@H](CCC(=O)O)N XOFYVODYSNKPDK-AVGNSLFASA-N 0.000 description 2
- DNPCBMNFQVTHMA-DCAQKATOSA-N Glu-Leu-Gln Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O DNPCBMNFQVTHMA-DCAQKATOSA-N 0.000 description 2
- YKBUCXNNBYZYAY-MNXVOIDGSA-N Glu-Lys-Ile Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O YKBUCXNNBYZYAY-MNXVOIDGSA-N 0.000 description 2
- AQNYKMCFCCZEEL-JYJNAYRXSA-N Glu-Lys-Tyr Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 AQNYKMCFCCZEEL-JYJNAYRXSA-N 0.000 description 2
- QNJNPKSWAHPYGI-JYJNAYRXSA-N Glu-Phe-Leu Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](CC(C)C)C(O)=O)CC1=CC=CC=C1 QNJNPKSWAHPYGI-JYJNAYRXSA-N 0.000 description 2
- AAJHGGDRKHYSDH-GUBZILKMSA-N Glu-Pro-Gln Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CCC(=O)O)N)C(=O)N[C@@H](CCC(=O)N)C(=O)O AAJHGGDRKHYSDH-GUBZILKMSA-N 0.000 description 2
- MLILEEIVMRUYBX-NHCYSSNCSA-N Glu-Val-Arg Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O MLILEEIVMRUYBX-NHCYSSNCSA-N 0.000 description 2
- BRFJMRSRMOMIMU-WHFBIAKZSA-N Gly-Ala-Asn Chemical compound NCC(=O)N[C@@H](C)C(=O)N[C@@H](CC(N)=O)C(O)=O BRFJMRSRMOMIMU-WHFBIAKZSA-N 0.000 description 2
- LJPIRKICOISLKN-WHFBIAKZSA-N Gly-Ala-Ser Chemical compound NCC(=O)N[C@@H](C)C(=O)N[C@@H](CO)C(O)=O LJPIRKICOISLKN-WHFBIAKZSA-N 0.000 description 2
- RJIVPOXLQFJRTG-LURJTMIESA-N Gly-Arg-Gly Chemical compound OC(=O)CNC(=O)[C@@H](NC(=O)CN)CCCN=C(N)N RJIVPOXLQFJRTG-LURJTMIESA-N 0.000 description 2
- WKJKBELXHCTHIJ-WPRPVWTQSA-N Gly-Arg-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)CN)CCCN=C(N)N WKJKBELXHCTHIJ-WPRPVWTQSA-N 0.000 description 2
- CIMULJZTTOBOPN-WHFBIAKZSA-N Gly-Asn-Asn Chemical compound NCC(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O CIMULJZTTOBOPN-WHFBIAKZSA-N 0.000 description 2
- GGEJHJIXRBTJPD-BYPYZUCNSA-N Gly-Asn-Gly Chemical compound NCC(=O)N[C@@H](CC(N)=O)C(=O)NCC(O)=O GGEJHJIXRBTJPD-BYPYZUCNSA-N 0.000 description 2
- FMNHBTKMRFVGRO-FOHZUACHSA-N Gly-Asn-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)[C@H](CC(N)=O)NC(=O)CN FMNHBTKMRFVGRO-FOHZUACHSA-N 0.000 description 2
- SABZDFAAOJATBR-QWRGUYRKSA-N Gly-Cys-Phe Chemical compound [H]NCC(=O)N[C@@H](CS)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O SABZDFAAOJATBR-QWRGUYRKSA-N 0.000 description 2
- BUEFQXUHTUZXHR-LURJTMIESA-N Gly-Gly-Pro zwitterion Chemical compound NCC(=O)NCC(=O)N1CCC[C@H]1C(O)=O BUEFQXUHTUZXHR-LURJTMIESA-N 0.000 description 2
- NNCSJUBVFBDDLC-YUMQZZPRSA-N Gly-Leu-Ser Chemical compound NCC(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O NNCSJUBVFBDDLC-YUMQZZPRSA-N 0.000 description 2
- OQQKUTVULYLCDG-ONGXEEELSA-N Gly-Lys-Val Chemical compound CC(C)[C@H](NC(=O)[C@H](CCCCN)NC(=O)CN)C(O)=O OQQKUTVULYLCDG-ONGXEEELSA-N 0.000 description 2
- OJNZVYSGVYLQIN-BQBZGAKWSA-N Gly-Met-Asp Chemical compound [H]NCC(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(O)=O)C(O)=O OJNZVYSGVYLQIN-BQBZGAKWSA-N 0.000 description 2
- HFPVRZWORNJRRC-UWVGGRQHSA-N Gly-Pro-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)CN HFPVRZWORNJRRC-UWVGGRQHSA-N 0.000 description 2
- SOEGEPHNZOISMT-BYPYZUCNSA-N Gly-Ser-Gly Chemical compound NCC(=O)N[C@@H](CO)C(=O)NCC(O)=O SOEGEPHNZOISMT-BYPYZUCNSA-N 0.000 description 2
- 235000010469 Glycine max Nutrition 0.000 description 2
- 244000068988 Glycine max Species 0.000 description 2
- VSLXGYMEHVAJBH-DLOVCJGASA-N His-Ala-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(O)=O VSLXGYMEHVAJBH-DLOVCJGASA-N 0.000 description 2
- FIMNVXRZGUAGBI-AVGNSLFASA-N His-Glu-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O FIMNVXRZGUAGBI-AVGNSLFASA-N 0.000 description 2
- JSHOVJTVPXJFTE-HOCLYGCPSA-N His-Gly-Trp Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)NCC(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(O)=O JSHOVJTVPXJFTE-HOCLYGCPSA-N 0.000 description 2
- MFQVZYSPCIZFMR-MGHWNKPDSA-N His-Ile-Phe Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H](CC2=CN=CN2)N MFQVZYSPCIZFMR-MGHWNKPDSA-N 0.000 description 2
- VZIFYHYNQDIPLI-HJWJTTGWSA-N Ile-Arg-Phe Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)N VZIFYHYNQDIPLI-HJWJTTGWSA-N 0.000 description 2
- CWJQMCPYXNVMBS-STECZYCISA-N Ile-Arg-Tyr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)O)N CWJQMCPYXNVMBS-STECZYCISA-N 0.000 description 2
- QIHJTGSVGIPHIW-QSFUFRPTSA-N Ile-Asn-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](C(C)C)C(=O)O)N QIHJTGSVGIPHIW-QSFUFRPTSA-N 0.000 description 2
- YBJWJQQBWRARLT-KBIXCLLPSA-N Ile-Gln-Ser Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(O)=O YBJWJQQBWRARLT-KBIXCLLPSA-N 0.000 description 2
- FTUZWJVSNZMLPI-RVMXOQNASA-N Ile-Met-Pro Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)N1CCC[C@@H]1C(=O)O)N FTUZWJVSNZMLPI-RVMXOQNASA-N 0.000 description 2
- FBGXMKUWQFPHFB-JBDRJPRFSA-N Ile-Ser-Cys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CS)C(=O)O)N FBGXMKUWQFPHFB-JBDRJPRFSA-N 0.000 description 2
- ZNOBVZFCHNHKHA-KBIXCLLPSA-N Ile-Ser-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N ZNOBVZFCHNHKHA-KBIXCLLPSA-N 0.000 description 2
- YBKKLDBBPFIXBQ-MBLNEYKQSA-N Ile-Thr-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)NCC(=O)O)N YBKKLDBBPFIXBQ-MBLNEYKQSA-N 0.000 description 2
- JZBVBOKASHNXAD-NAKRPEOUSA-N Ile-Val-Ser Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CO)C(=O)O)N JZBVBOKASHNXAD-NAKRPEOUSA-N 0.000 description 2
- RCFDOSNHHZGBOY-UHFFFAOYSA-N L-isoleucyl-L-alanine Natural products CCC(C)C(N)C(=O)NC(C)C(O)=O RCFDOSNHHZGBOY-UHFFFAOYSA-N 0.000 description 2
- LHSGPCFBGJHPCY-UHFFFAOYSA-N L-leucine-L-tyrosine Natural products CC(C)CC(N)C(=O)NC(C(O)=O)CC1=CC=C(O)C=C1 LHSGPCFBGJHPCY-UHFFFAOYSA-N 0.000 description 2
- 108091026898 Leader sequence (mRNA) Proteins 0.000 description 2
- BPANDPNDMJHFEV-CIUDSAMLSA-N Leu-Asp-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C)C(O)=O BPANDPNDMJHFEV-CIUDSAMLSA-N 0.000 description 2
- ILJREDZFPHTUIE-GUBZILKMSA-N Leu-Asp-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O ILJREDZFPHTUIE-GUBZILKMSA-N 0.000 description 2
- DLCOFDAHNMMQPP-SRVKXCTJSA-N Leu-Asp-Leu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O DLCOFDAHNMMQPP-SRVKXCTJSA-N 0.000 description 2
- XVSJMWYYLHPDKY-DCAQKATOSA-N Leu-Asp-Met Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCSC)C(O)=O XVSJMWYYLHPDKY-DCAQKATOSA-N 0.000 description 2
- KAFOIVJDVSZUMD-UHFFFAOYSA-N Leu-Gln-Gln Natural products CC(C)CC(N)C(=O)NC(CCC(N)=O)C(=O)NC(CCC(N)=O)C(O)=O KAFOIVJDVSZUMD-UHFFFAOYSA-N 0.000 description 2
- BOFAFKVZQUMTID-AVGNSLFASA-N Leu-Gln-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N BOFAFKVZQUMTID-AVGNSLFASA-N 0.000 description 2
- NEEOBPIXKWSBRF-IUCAKERBSA-N Leu-Glu-Gly Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)NCC(O)=O NEEOBPIXKWSBRF-IUCAKERBSA-N 0.000 description 2
- LLBQJYDYOLIQAI-JYJNAYRXSA-N Leu-Glu-Tyr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O LLBQJYDYOLIQAI-JYJNAYRXSA-N 0.000 description 2
- HYMLKESRWLZDBR-WEDXCCLWSA-N Leu-Gly-Thr Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(O)=O HYMLKESRWLZDBR-WEDXCCLWSA-N 0.000 description 2
- JKSIBWITFMQTOA-XUXIUFHCSA-N Leu-Ile-Val Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C(C)C)C(O)=O JKSIBWITFMQTOA-XUXIUFHCSA-N 0.000 description 2
- JNDYEOUZBLOVOF-AVGNSLFASA-N Leu-Leu-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O JNDYEOUZBLOVOF-AVGNSLFASA-N 0.000 description 2
- YOKVEHGYYQEQOP-QWRGUYRKSA-N Leu-Leu-Gly Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O YOKVEHGYYQEQOP-QWRGUYRKSA-N 0.000 description 2
- RXGLHDWAZQECBI-SRVKXCTJSA-N Leu-Leu-Ser Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O RXGLHDWAZQECBI-SRVKXCTJSA-N 0.000 description 2
- HVHRPWQEQHIQJF-AVGNSLFASA-N Leu-Lys-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O HVHRPWQEQHIQJF-AVGNSLFASA-N 0.000 description 2
- GCXGCIYIHXSKAY-ULQDDVLXSA-N Leu-Phe-Arg Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O GCXGCIYIHXSKAY-ULQDDVLXSA-N 0.000 description 2
- KQFZKDITNUEVFJ-JYJNAYRXSA-N Leu-Phe-Gln Chemical compound NC(=O)CC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)CC(C)C)CC1=CC=CC=C1 KQFZKDITNUEVFJ-JYJNAYRXSA-N 0.000 description 2
- INCJJHQRZGQLFC-KBPBESRZSA-N Leu-Phe-Gly Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)NCC(O)=O INCJJHQRZGQLFC-KBPBESRZSA-N 0.000 description 2
- PJWOOBTYQNNRBF-BZSNNMDCSA-N Leu-Phe-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCCCN)C(=O)O)N PJWOOBTYQNNRBF-BZSNNMDCSA-N 0.000 description 2
- XXXXOVFBXRERQL-ULQDDVLXSA-N Leu-Pro-Phe Chemical compound CC(C)C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 XXXXOVFBXRERQL-ULQDDVLXSA-N 0.000 description 2
- JIHDFWWRYHSAQB-GUBZILKMSA-N Leu-Ser-Glu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCC(O)=O JIHDFWWRYHSAQB-GUBZILKMSA-N 0.000 description 2
- MVJRBCJCRYGCKV-GVXVVHGQSA-N Leu-Val-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O MVJRBCJCRYGCKV-GVXVVHGQSA-N 0.000 description 2
- 235000007688 Lycopersicon esculentum Nutrition 0.000 description 2
- IXHKPDJKKCUKHS-GARJFASQSA-N Lys-Ala-Pro Chemical compound C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CCCCN)N IXHKPDJKKCUKHS-GARJFASQSA-N 0.000 description 2
- FLCMXEFCTLXBTL-DCAQKATOSA-N Lys-Asp-Arg Chemical compound C(CCN)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)N FLCMXEFCTLXBTL-DCAQKATOSA-N 0.000 description 2
- GJJQCBVRWDGLMQ-GUBZILKMSA-N Lys-Glu-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O GJJQCBVRWDGLMQ-GUBZILKMSA-N 0.000 description 2
- LPAJOCKCPRZEAG-MNXVOIDGSA-N Lys-Glu-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)[C@H](CCC(O)=O)NC(=O)[C@@H](N)CCCCN LPAJOCKCPRZEAG-MNXVOIDGSA-N 0.000 description 2
- ZXFRGTAIIZHNHG-AJNGGQMLSA-N Lys-Ile-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)O)NC(=O)[C@H](CCCCN)N ZXFRGTAIIZHNHG-AJNGGQMLSA-N 0.000 description 2
- RBEATVHTWHTHTJ-KKUMJFAQSA-N Lys-Leu-Lys Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(O)=O RBEATVHTWHTHTJ-KKUMJFAQSA-N 0.000 description 2
- XOQMURBBIXRRCR-SRVKXCTJSA-N Lys-Lys-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CCCCN XOQMURBBIXRRCR-SRVKXCTJSA-N 0.000 description 2
- GAHJXEMYXKLZRQ-AJNGGQMLSA-N Lys-Lys-Ile Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O GAHJXEMYXKLZRQ-AJNGGQMLSA-N 0.000 description 2
- YDDDRTIPNTWGIG-SRVKXCTJSA-N Lys-Lys-Ser Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(O)=O YDDDRTIPNTWGIG-SRVKXCTJSA-N 0.000 description 2
- 240000004658 Medicago sativa Species 0.000 description 2
- 235000017587 Medicago sativa ssp. sativa Nutrition 0.000 description 2
- ULNXMMYXQKGNPG-LPEHRKFASA-N Met-Ala-Pro Chemical compound C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CCSC)N ULNXMMYXQKGNPG-LPEHRKFASA-N 0.000 description 2
- YCUSPBPZVJDMII-YUMQZZPRSA-N Met-Gly-Glu Chemical compound CSCC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCC(O)=O YCUSPBPZVJDMII-YUMQZZPRSA-N 0.000 description 2
- LRALLISKBZNSKN-BQBZGAKWSA-N Met-Gly-Ser Chemical compound CSCC[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O LRALLISKBZNSKN-BQBZGAKWSA-N 0.000 description 2
- AFFKUNVPPLQUGA-DCAQKATOSA-N Met-Leu-Ala Chemical compound [H]N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](C)C(O)=O AFFKUNVPPLQUGA-DCAQKATOSA-N 0.000 description 2
- DBXMFHGGHMXYHY-DCAQKATOSA-N Met-Leu-Ser Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O DBXMFHGGHMXYHY-DCAQKATOSA-N 0.000 description 2
- JOYFULUKJRJCSX-IUCAKERBSA-N Met-Met-Gly Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCSC)C(=O)NCC(O)=O JOYFULUKJRJCSX-IUCAKERBSA-N 0.000 description 2
- KZNQNBZMBZJQJO-UHFFFAOYSA-N N-glycyl-L-proline Natural products NCC(=O)N1CCCC1C(O)=O KZNQNBZMBZJQJO-UHFFFAOYSA-N 0.000 description 2
- 108010079364 N-glycylalanine Proteins 0.000 description 2
- 108010087066 N2-tryptophyllysine Proteins 0.000 description 2
- 101100342977 Neurospora crassa (strain ATCC 24698 / 74-OR23-1A / CBS 708.71 / DSM 1257 / FGSC 987) leu-1 gene Proteins 0.000 description 2
- VJLLEKDQJSMHRU-STQMWFEESA-N Phe-Gly-Met Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)NCC(=O)N[C@@H](CCSC)C(O)=O VJLLEKDQJSMHRU-STQMWFEESA-N 0.000 description 2
- YVXPUUOTMVBKDO-IHRRRGAJSA-N Phe-Pro-Cys Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CC2=CC=CC=C2)N)C(=O)N[C@@H](CS)C(=O)O YVXPUUOTMVBKDO-IHRRRGAJSA-N 0.000 description 2
- GMWNQSGWWGKTSF-LFSVMHDDSA-N Phe-Thr-Ala Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C)C(O)=O GMWNQSGWWGKTSF-LFSVMHDDSA-N 0.000 description 2
- MSSXKZBDKZAHCX-UNQGMJICSA-N Phe-Thr-Val Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C(C)C)C(O)=O MSSXKZBDKZAHCX-UNQGMJICSA-N 0.000 description 2
- 241000219000 Populus Species 0.000 description 2
- SKICPQLTOXGWGO-GARJFASQSA-N Pro-Gln-Pro Chemical compound C1C[C@H](NC1)C(=O)N[C@@H](CCC(=O)N)C(=O)N2CCC[C@@H]2C(=O)O SKICPQLTOXGWGO-GARJFASQSA-N 0.000 description 2
- YSUZKYSRAFNLRB-ULQDDVLXSA-N Pro-Gln-Trp Chemical compound N([C@@H](CCC(=O)N)C(=O)N[C@@H](CC=1C2=CC=CC=C2NC=1)C(O)=O)C(=O)[C@@H]1CCCN1 YSUZKYSRAFNLRB-ULQDDVLXSA-N 0.000 description 2
- RPLMFKUKFZOTER-AVGNSLFASA-N Pro-Met-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CCSC)NC(=O)[C@@H]1CCCN1 RPLMFKUKFZOTER-AVGNSLFASA-N 0.000 description 2
- FYKUEXMZYFIZKA-DCAQKATOSA-N Pro-Pro-Gln Chemical compound [H]N1CCC[C@H]1C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCC(N)=O)C(O)=O FYKUEXMZYFIZKA-DCAQKATOSA-N 0.000 description 2
- CGSOWZUPLOKYOR-AVGNSLFASA-N Pro-Pro-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@H]1NCCC1 CGSOWZUPLOKYOR-AVGNSLFASA-N 0.000 description 2
- LZHHZYDPMZEMRX-STQMWFEESA-N Pro-Tyr-Gly Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)NCC(O)=O LZHHZYDPMZEMRX-STQMWFEESA-N 0.000 description 2
- ZMLRZBWCXPQADC-TUAOUCFPSA-N Pro-Val-Pro Chemical compound CC(C)[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@@H]2CCCN2 ZMLRZBWCXPQADC-TUAOUCFPSA-N 0.000 description 2
- VGNYHOBZJKWRGI-CIUDSAMLSA-N Ser-Asn-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CC(N)=O)NC(=O)[C@@H](N)CO VGNYHOBZJKWRGI-CIUDSAMLSA-N 0.000 description 2
- ZOHGLPQGEHSLPD-FXQIFTODSA-N Ser-Gln-Glu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O ZOHGLPQGEHSLPD-FXQIFTODSA-N 0.000 description 2
- IAORETPTUDBBGV-CIUDSAMLSA-N Ser-Leu-Cys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CO)N IAORETPTUDBBGV-CIUDSAMLSA-N 0.000 description 2
- YUJLIIRMIAGMCQ-CIUDSAMLSA-N Ser-Leu-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O YUJLIIRMIAGMCQ-CIUDSAMLSA-N 0.000 description 2
- JWOBLHJRDADHLN-KKUMJFAQSA-N Ser-Leu-Tyr Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O JWOBLHJRDADHLN-KKUMJFAQSA-N 0.000 description 2
- VIIJCAQMJBHSJH-FXQIFTODSA-N Ser-Met-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CO)C(O)=O VIIJCAQMJBHSJH-FXQIFTODSA-N 0.000 description 2
- NQZFFLBPNDLTPO-DLOVCJGASA-N Ser-Phe-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CC1=CC=CC=C1)NC(=O)[C@H](CO)N NQZFFLBPNDLTPO-DLOVCJGASA-N 0.000 description 2
- ADJDNJCSPNFFPI-FXQIFTODSA-N Ser-Pro-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)CO ADJDNJCSPNFFPI-FXQIFTODSA-N 0.000 description 2
- HHJFMHQYEAAOBM-ZLUOBGJFSA-N Ser-Ser-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H](C)C(O)=O HHJFMHQYEAAOBM-ZLUOBGJFSA-N 0.000 description 2
- KQNDIKOYWZTZIX-FXQIFTODSA-N Ser-Ser-Arg Chemical compound OC[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCCNC(N)=N KQNDIKOYWZTZIX-FXQIFTODSA-N 0.000 description 2
- NADLKBTYNKUJEP-KATARQTJSA-N Ser-Thr-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(O)=O NADLKBTYNKUJEP-KATARQTJSA-N 0.000 description 2
- SNXUIBACCONSOH-BWBBJGPYSA-N Ser-Thr-Ser Chemical compound OC[C@H](N)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@@H](CO)C(O)=O SNXUIBACCONSOH-BWBBJGPYSA-N 0.000 description 2
- WMZVVNLPHFSUPA-BPUTZDHNSA-N Ser-Trp-Arg Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](CO)N)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O)=CNC2=C1 WMZVVNLPHFSUPA-BPUTZDHNSA-N 0.000 description 2
- 240000003768 Solanum lycopersicum Species 0.000 description 2
- 244000061458 Solanum melongena Species 0.000 description 2
- IGROJMCBGRFRGI-YTLHQDLWSA-N Thr-Ala-Ala Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@@H](C)C(O)=O IGROJMCBGRFRGI-YTLHQDLWSA-N 0.000 description 2
- YBXMGKCLOPDEKA-NUMRIWBASA-N Thr-Asp-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O YBXMGKCLOPDEKA-NUMRIWBASA-N 0.000 description 2
- SHOMROOOQBDGRL-JHEQGTHGSA-N Thr-Glu-Gly Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(=O)NCC(O)=O SHOMROOOQBDGRL-JHEQGTHGSA-N 0.000 description 2
- DJDSEDOKJTZBAR-ZDLURKLDSA-N Thr-Gly-Ser Chemical compound C[C@@H](O)[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O DJDSEDOKJTZBAR-ZDLURKLDSA-N 0.000 description 2
- XSTGOZBBXFKGHA-YJRXYDGGSA-N Thr-His-His Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)N)O XSTGOZBBXFKGHA-YJRXYDGGSA-N 0.000 description 2
- UYTYTDMCDBPDSC-URLPEUOOSA-N Thr-Ile-Phe Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H]([C@@H](C)O)N UYTYTDMCDBPDSC-URLPEUOOSA-N 0.000 description 2
- RRRRCRYTLZVCEN-HJGDQZAQSA-N Thr-Leu-Asp Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O RRRRCRYTLZVCEN-HJGDQZAQSA-N 0.000 description 2
- LKJCABTUFGTPPY-HJGDQZAQSA-N Thr-Pro-Gln Chemical compound C[C@@H](O)[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCC(N)=O)C(O)=O LKJCABTUFGTPPY-HJGDQZAQSA-N 0.000 description 2
- SGAOHNPSEPVAFP-ZDLURKLDSA-N Thr-Ser-Gly Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(=O)NCC(O)=O SGAOHNPSEPVAFP-ZDLURKLDSA-N 0.000 description 2
- IEZVHOULSUULHD-XGEHTFHBSA-N Thr-Ser-Val Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(O)=O IEZVHOULSUULHD-XGEHTFHBSA-N 0.000 description 2
- 108091036066 Three prime untranslated region Proteins 0.000 description 2
- 241000062949 Tremula Species 0.000 description 2
- QNTBGBCOEYNAPV-CWRNSKLLSA-N Trp-Asn-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC(=O)N)NC(=O)[C@H](CC2=CNC3=CC=CC=C32)N)C(=O)O QNTBGBCOEYNAPV-CWRNSKLLSA-N 0.000 description 2
- VKMOGXREKGVZAF-QEJZJMRPSA-N Trp-Asp-Gln Chemical compound C1=CC=C2C(=C1)C(=CN2)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N VKMOGXREKGVZAF-QEJZJMRPSA-N 0.000 description 2
- MWHOLXNKRKRQQH-XIRDDKMYSA-N Trp-Asp-His Chemical compound C1=CC=C2C(=C1)C(=CN2)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CC3=CN=CN3)C(=O)O)N MWHOLXNKRKRQQH-XIRDDKMYSA-N 0.000 description 2
- GTNCSPKYWCJZAC-XIRDDKMYSA-N Trp-Asp-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H](CC1=CNC2=CC=CC=C21)N GTNCSPKYWCJZAC-XIRDDKMYSA-N 0.000 description 2
- YXONONCLMLHWJX-SZMVWBNQSA-N Trp-Glu-Leu Chemical compound C1=CC=C2C(C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O)=CNC2=C1 YXONONCLMLHWJX-SZMVWBNQSA-N 0.000 description 2
- VFJIWSJKZJTQII-SRVKXCTJSA-N Tyr-Asp-Ser Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(O)=O VFJIWSJKZJTQII-SRVKXCTJSA-N 0.000 description 2
- HVHJYXDXRIWELT-RYUDHWBXSA-N Tyr-Glu-Gly Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(O)=O)C(=O)NCC(O)=O HVHJYXDXRIWELT-RYUDHWBXSA-N 0.000 description 2
- VTCKHZJKWQENKX-KBPBESRZSA-N Tyr-Lys-Gly Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCCCN)C(=O)NCC(O)=O VTCKHZJKWQENKX-KBPBESRZSA-N 0.000 description 2
- XYBNMHRFAUKPAW-IHRRRGAJSA-N Tyr-Ser-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CC1=CC=C(C=C1)O)N XYBNMHRFAUKPAW-IHRRRGAJSA-N 0.000 description 2
- LTFLDDDGWOVIHY-NAKRPEOUSA-N Val-Ala-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](C)NC(=O)[C@H](C(C)C)N LTFLDDDGWOVIHY-NAKRPEOUSA-N 0.000 description 2
- ZLFHAAGHGQBQQN-AEJSXWLSSA-N Val-Ala-Pro Chemical compound C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](C(C)C)N ZLFHAAGHGQBQQN-AEJSXWLSSA-N 0.000 description 2
- ZLFHAAGHGQBQQN-GUBZILKMSA-N Val-Ala-Pro Natural products CC(C)[C@H](N)C(=O)N[C@@H](C)C(=O)N1CCC[C@H]1C(O)=O ZLFHAAGHGQBQQN-GUBZILKMSA-N 0.000 description 2
- LIQJSDDOULTANC-QSFUFRPTSA-N Val-Asn-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](C(C)C)N LIQJSDDOULTANC-QSFUFRPTSA-N 0.000 description 2
- FXVDGDZRYLFQKY-WPRPVWTQSA-N Val-Gly-Met Chemical compound CSCC[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)C(C)C FXVDGDZRYLFQKY-WPRPVWTQSA-N 0.000 description 2
- NZGOVKLVQNOEKP-YDHLFZDLSA-N Val-Phe-Asn Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(=O)N)C(=O)O)N NZGOVKLVQNOEKP-YDHLFZDLSA-N 0.000 description 2
- KSFXWENSJABBFI-ZKWXMUAHSA-N Val-Ser-Asn Chemical compound [H]N[C@@H](C(C)C)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(N)=O)C(O)=O KSFXWENSJABBFI-ZKWXMUAHSA-N 0.000 description 2
- UFCHCOKFAGOQSF-BQFCYCMXSA-N Val-Trp-Glu Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N UFCHCOKFAGOQSF-BQFCYCMXSA-N 0.000 description 2
- 230000001133 acceleration Effects 0.000 description 2
- 108010008685 alanyl-glutamyl-aspartic acid Proteins 0.000 description 2
- 108010076324 alanyl-glycyl-glycine Proteins 0.000 description 2
- 235000001014 amino acid Nutrition 0.000 description 2
- 230000019552 anatomical structure morphogenesis Effects 0.000 description 2
- 108010043240 arginyl-leucyl-glycine Proteins 0.000 description 2
- 108010068380 arginylarginine Proteins 0.000 description 2
- 108010047857 aspartylglycine Proteins 0.000 description 2
- 241001233866 asterids Species 0.000 description 2
- 230000001580 bacterial effect Effects 0.000 description 2
- 238000010367 cloning Methods 0.000 description 2
- 244000038559 crop plants Species 0.000 description 2
- 230000003111 delayed effect Effects 0.000 description 2
- FSXRLASFHBWESK-UHFFFAOYSA-N dipeptide phenylalanyl-tyrosine Natural products C=1C=C(O)C=CC=1CC(C(O)=O)NC(=O)C(N)CC1=CC=CC=C1 FSXRLASFHBWESK-UHFFFAOYSA-N 0.000 description 2
- 241001233957 eudicotyledons Species 0.000 description 2
- 238000010353 genetic engineering Methods 0.000 description 2
- 108010079547 glutamylmethionine Proteins 0.000 description 2
- 108010019832 glycyl-asparaginyl-glycine Proteins 0.000 description 2
- 108010051307 glycyl-glycyl-proline Proteins 0.000 description 2
- 108010078326 glycyl-glycyl-valine Proteins 0.000 description 2
- 108010089804 glycyl-threonine Proteins 0.000 description 2
- 108010050848 glycylleucine Proteins 0.000 description 2
- 108010077515 glycylproline Proteins 0.000 description 2
- 108010040030 histidinoalanine Proteins 0.000 description 2
- 239000005556 hormone Substances 0.000 description 2
- 229940088597 hormone Drugs 0.000 description 2
- 238000000338 in vitro Methods 0.000 description 2
- 230000008595 infiltration Effects 0.000 description 2
- 238000001764 infiltration Methods 0.000 description 2
- 230000000366 juvenile effect Effects 0.000 description 2
- 108010044311 leucyl-glycyl-glycine Proteins 0.000 description 2
- 108010012058 leucyltyrosine Proteins 0.000 description 2
- 108010054155 lysyllysine Proteins 0.000 description 2
- 230000001404 mediated effect Effects 0.000 description 2
- 231100001222 nononcogenic Toxicity 0.000 description 2
- 239000002245 particle Substances 0.000 description 2
- 108010024607 phenylalanylalanine Proteins 0.000 description 2
- 108010025488 pinealon Proteins 0.000 description 2
- 230000008488 polyadenylation Effects 0.000 description 2
- 102000040430 polynucleotide Human genes 0.000 description 2
- 108091033319 polynucleotide Proteins 0.000 description 2
- 239000002157 polynucleotide Substances 0.000 description 2
- 239000001054 red pigment Substances 0.000 description 2
- 230000022532 regulation of transcription, DNA-dependent Effects 0.000 description 2
- 239000006152 selective media Substances 0.000 description 2
- 239000002689 soil Substances 0.000 description 2
- 108010061238 threonyl-glycine Proteins 0.000 description 2
- 108010003137 tyrosyltyrosine Proteins 0.000 description 2
- 108010015385 valyl-prolyl-proline Proteins 0.000 description 2
- 230000003612 virological effect Effects 0.000 description 2
- DIGQNXIGRZPYDK-WKSCXVIASA-N (2R)-6-amino-2-[[2-[[(2S)-2-[[2-[[(2R)-2-[[(2S)-2-[[(2R,3S)-2-[[2-[[(2S)-2-[[2-[[(2S)-2-[[(2S)-2-[[(2R)-2-[[(2S,3S)-2-[[(2R)-2-[[(2S)-2-[[(2S)-2-[[(2S)-2-[[2-[[(2S)-2-[[(2R)-2-[[2-[[2-[[2-[(2-amino-1-hydroxyethylidene)amino]-3-carboxy-1-hydroxypropylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1-hydroxyethylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1,3-dihydroxypropylidene]amino]-1-hydroxyethylidene]amino]-1-hydroxypropylidene]amino]-1,3-dihydroxypropylidene]amino]-1,3-dihydroxypropylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1,3-dihydroxybutylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1-hydroxypropylidene]amino]-1,3-dihydroxypropylidene]amino]-1-hydroxyethylidene]amino]-1,5-dihydroxy-5-iminopentylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1,3-dihydroxybutylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1,3-dihydroxypropylidene]amino]-1-hydroxyethylidene]amino]-1-hydroxy-3-sulfanylpropylidene]amino]-1-hydroxyethylidene]amino]hexanoic acid Chemical compound C[C@@H]([C@@H](C(=N[C@@H](CS)C(=N[C@@H](C)C(=N[C@@H](CO)C(=NCC(=N[C@@H](CCC(=N)O)C(=NC(CS)C(=N[C@H]([C@H](C)O)C(=N[C@H](CS)C(=N[C@H](CO)C(=NCC(=N[C@H](CS)C(=NCC(=N[C@H](CCCCN)C(=O)O)O)O)O)O)O)O)O)O)O)O)O)O)O)N=C([C@H](CS)N=C([C@H](CO)N=C([C@H](CO)N=C([C@H](C)N=C(CN=C([C@H](CO)N=C([C@H](CS)N=C(CN=C(C(CS)N=C(C(CC(=O)O)N=C(CN)O)O)O)O)O)O)O)O)O)O)O)O DIGQNXIGRZPYDK-WKSCXVIASA-N 0.000 description 1
- CWFMWBHMIMNZLN-NAKRPEOUSA-N (2s)-1-[(2s)-2-[[(2s,3s)-2-amino-3-methylpentanoyl]amino]propanoyl]pyrrolidine-2-carboxylic acid Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](C)C(=O)N1CCC[C@H]1C(O)=O CWFMWBHMIMNZLN-NAKRPEOUSA-N 0.000 description 1
- 101710169336 5'-deoxyadenosine deaminase Proteins 0.000 description 1
- 102000055025 Adenosine deaminases Human genes 0.000 description 1
- OMMDTNGURYRDAC-NRPADANISA-N Ala-Glu-Val Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O OMMDTNGURYRDAC-NRPADANISA-N 0.000 description 1
- PCIFXPRIFWKWLK-YUMQZZPRSA-N Ala-Gly-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)CNC(=O)[C@H](C)N PCIFXPRIFWKWLK-YUMQZZPRSA-N 0.000 description 1
- GRPHQEMIFDPKOE-HGNGGELXSA-N Ala-His-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CCC(O)=O)C(O)=O GRPHQEMIFDPKOE-HGNGGELXSA-N 0.000 description 1
- GSHKMNKPMLXSQW-KBIXCLLPSA-N Ala-Ile-Gln Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)O)NC(=O)[C@H](C)N GSHKMNKPMLXSQW-KBIXCLLPSA-N 0.000 description 1
- XHNLCGXYBXNRIS-BJDJZHNGSA-N Ala-Lys-Ile Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O XHNLCGXYBXNRIS-BJDJZHNGSA-N 0.000 description 1
- NLOMBWNGESDVJU-GUBZILKMSA-N Ala-Met-Arg Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O NLOMBWNGESDVJU-GUBZILKMSA-N 0.000 description 1
- AWNAEZICPNGAJK-FXQIFTODSA-N Ala-Met-Ser Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CO)C(O)=O AWNAEZICPNGAJK-FXQIFTODSA-N 0.000 description 1
- 244000291564 Allium cepa Species 0.000 description 1
- 235000002732 Allium cepa var. cepa Nutrition 0.000 description 1
- KWKQGHSSNHPGOW-BQBZGAKWSA-N Arg-Ala-Gly Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](C)C(=O)NCC(O)=O KWKQGHSSNHPGOW-BQBZGAKWSA-N 0.000 description 1
- XUBLMYHWSFRACH-CYDGBPFRSA-N Arg-Asn-Gln-Arg Chemical compound NC(=N)NCCC[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O XUBLMYHWSFRACH-CYDGBPFRSA-N 0.000 description 1
- UZGFHWIJWPUPOH-IHRRRGAJSA-N Arg-Leu-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N UZGFHWIJWPUPOH-IHRRRGAJSA-N 0.000 description 1
- AIFHRTPABBBHKU-RCWTZXSCSA-N Arg-Thr-Arg Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O AIFHRTPABBBHKU-RCWTZXSCSA-N 0.000 description 1
- NPDLYUOYAGBHFB-WDSKDSINSA-N Asn-Arg Chemical compound NC(=O)C[C@H](N)C(=O)N[C@H](C(O)=O)CCCN=C(N)N NPDLYUOYAGBHFB-WDSKDSINSA-N 0.000 description 1
- CIBWFJFMOBIFTE-CIUDSAMLSA-N Asn-Arg-Gln Chemical compound C(C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)O)NC(=O)[C@H](CC(=O)N)N)CN=C(N)N CIBWFJFMOBIFTE-CIUDSAMLSA-N 0.000 description 1
- UGXVKHRDGLYFKR-CIUDSAMLSA-N Asn-Asp-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(O)=O)NC(=O)[C@@H](N)CC(N)=O UGXVKHRDGLYFKR-CIUDSAMLSA-N 0.000 description 1
- HLTLEIXYIJDFOY-ZLUOBGJFSA-N Asn-Cys-Asn Chemical compound NC(=O)C[C@H](N)C(=O)N[C@@H](CS)C(=O)N[C@@H](CC(N)=O)C(O)=O HLTLEIXYIJDFOY-ZLUOBGJFSA-N 0.000 description 1
- WPOLSNAQGVHROR-GUBZILKMSA-N Asn-Gln-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC(=O)N)N WPOLSNAQGVHROR-GUBZILKMSA-N 0.000 description 1
- WIDVAWAQBRAKTI-YUMQZZPRSA-N Asn-Leu-Gly Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O WIDVAWAQBRAKTI-YUMQZZPRSA-N 0.000 description 1
- MYVBTYXSWILFCG-BQBZGAKWSA-N Asn-Met-Gly Chemical compound CSCC[C@@H](C(=O)NCC(=O)O)NC(=O)[C@H](CC(=O)N)N MYVBTYXSWILFCG-BQBZGAKWSA-N 0.000 description 1
- MLJZMGIXXMTEPO-UBHSHLNASA-N Asn-Trp-Ser Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CO)C(O)=O MLJZMGIXXMTEPO-UBHSHLNASA-N 0.000 description 1
- HPNDBHLITCHRSO-WHFBIAKZSA-N Asp-Ala-Gly Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](C)C(=O)NCC(O)=O HPNDBHLITCHRSO-WHFBIAKZSA-N 0.000 description 1
- XLILXFRAKOYEJX-GUBZILKMSA-N Asp-Leu-Gln Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O XLILXFRAKOYEJX-GUBZILKMSA-N 0.000 description 1
- DWOGMPWRQQWPPF-GUBZILKMSA-N Asp-Leu-Glu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O DWOGMPWRQQWPPF-GUBZILKMSA-N 0.000 description 1
- LGGHQRZIJSYRHA-GUBZILKMSA-N Asp-Pro-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@@H]1CCCN1C(=O)[C@H](CC(=O)O)N LGGHQRZIJSYRHA-GUBZILKMSA-N 0.000 description 1
- FIAKNCXQFFKSSI-ZLUOBGJFSA-N Asp-Ser-Cys Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CS)C(O)=O FIAKNCXQFFKSSI-ZLUOBGJFSA-N 0.000 description 1
- 244000003416 Asparagus officinalis Species 0.000 description 1
- 235000005340 Asparagus officinalis Nutrition 0.000 description 1
- 229930192334 Auxin Natural products 0.000 description 1
- 235000007319 Avena orientalis Nutrition 0.000 description 1
- 244000075850 Avena orientalis Species 0.000 description 1
- 241000894006 Bacteria Species 0.000 description 1
- 235000016068 Berberis vulgaris Nutrition 0.000 description 1
- 241000335053 Beta vulgaris Species 0.000 description 1
- 235000011303 Brassica alboglabra Nutrition 0.000 description 1
- 235000006008 Brassica napus var napus Nutrition 0.000 description 1
- 235000011302 Brassica oleracea Nutrition 0.000 description 1
- 235000004221 Brassica oleracea var gemmifera Nutrition 0.000 description 1
- 235000012905 Brassica oleracea var viridis Nutrition 0.000 description 1
- 244000308368 Brassica oleracea var. gemmifera Species 0.000 description 1
- 235000010149 Brassica rapa subsp chinensis Nutrition 0.000 description 1
- 235000000536 Brassica rapa subsp pekinensis Nutrition 0.000 description 1
- 241000499436 Brassica rapa subsp. pekinensis Species 0.000 description 1
- 235000004977 Brassica sinapistrum Nutrition 0.000 description 1
- 241000218980 Brassicales Species 0.000 description 1
- 101100268548 Caenorhabditis elegans apl-1 gene Proteins 0.000 description 1
- 235000002566 Capsicum Nutrition 0.000 description 1
- 235000002568 Capsicum frutescens Nutrition 0.000 description 1
- 101710132601 Capsid protein Proteins 0.000 description 1
- 101710094648 Coat protein Proteins 0.000 description 1
- 108091026890 Coding region Proteins 0.000 description 1
- 229920000742 Cotton Polymers 0.000 description 1
- JPVYNHNXODAKFH-UHFFFAOYSA-N Cu2+ Chemical compound [Cu+2] JPVYNHNXODAKFH-UHFFFAOYSA-N 0.000 description 1
- 235000015510 Cucumis melo subsp melo Nutrition 0.000 description 1
- 235000009847 Cucumis melo var cantalupensis Nutrition 0.000 description 1
- 240000008067 Cucumis sativus Species 0.000 description 1
- 235000010799 Cucumis sativus var sativus Nutrition 0.000 description 1
- 235000009854 Cucurbita moschata Nutrition 0.000 description 1
- 240000001980 Cucurbita pepo Species 0.000 description 1
- 235000009852 Cucurbita pepo Nutrition 0.000 description 1
- UIKLEGZPIOXFHJ-DLOVCJGASA-N Cys-Phe-Ala Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](C)C(O)=O UIKLEGZPIOXFHJ-DLOVCJGASA-N 0.000 description 1
- 235000002767 Daucus carota Nutrition 0.000 description 1
- 244000000626 Daucus carota Species 0.000 description 1
- 101100364969 Dictyostelium discoideum scai gene Proteins 0.000 description 1
- 108090000790 Enzymes Proteins 0.000 description 1
- 102000004190 Enzymes Human genes 0.000 description 1
- 241000588724 Escherichia coli Species 0.000 description 1
- 101001091269 Escherichia coli Hygromycin-B 4-O-kinase Proteins 0.000 description 1
- IGNGBUVODQLMRJ-CIUDSAMLSA-N Gln-Ala-Met Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C)C(=O)N[C@@H](CCSC)C(O)=O IGNGBUVODQLMRJ-CIUDSAMLSA-N 0.000 description 1
- RGRMOYQUIJVQQD-SRVKXCTJSA-N Gln-Arg-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@H](CCC(=O)N)N RGRMOYQUIJVQQD-SRVKXCTJSA-N 0.000 description 1
- RMOCFPBLHAOTDU-ACZMJKKPSA-N Gln-Asn-Ser Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(O)=O RMOCFPBLHAOTDU-ACZMJKKPSA-N 0.000 description 1
- GPISLLFQNHELLK-DCAQKATOSA-N Gln-Gln-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CCC(=O)N)N GPISLLFQNHELLK-DCAQKATOSA-N 0.000 description 1
- XSBGUANSZDGULP-IUCAKERBSA-N Gln-Gly-Lys Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)NCC(=O)N[C@@H](CCCCN)C(O)=O XSBGUANSZDGULP-IUCAKERBSA-N 0.000 description 1
- GQZDDFRXSDGUNG-YVNDNENWSA-N Gln-Ile-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(N)=O)C(O)=O GQZDDFRXSDGUNG-YVNDNENWSA-N 0.000 description 1
- QBLMTCRYYTVUQY-GUBZILKMSA-N Gln-Leu-Asp Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O QBLMTCRYYTVUQY-GUBZILKMSA-N 0.000 description 1
- KHNJVFYHIKLUPD-SRVKXCTJSA-N Gln-Leu-Met Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCSC)C(=O)O)NC(=O)[C@H](CCC(=O)N)N KHNJVFYHIKLUPD-SRVKXCTJSA-N 0.000 description 1
- VNTGPISAOMAXRK-CIUDSAMLSA-N Gln-Pro-Ser Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O VNTGPISAOMAXRK-CIUDSAMLSA-N 0.000 description 1
- CVPXINNKRTZBMO-CIUDSAMLSA-N Glu-Arg-Asn Chemical compound C(C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CCC(=O)O)N)CN=C(N)N CVPXINNKRTZBMO-CIUDSAMLSA-N 0.000 description 1
- JPHYJQHPILOKHC-ACZMJKKPSA-N Glu-Asp-Asp Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O JPHYJQHPILOKHC-ACZMJKKPSA-N 0.000 description 1
- JVSBYEDSSRZQGV-GUBZILKMSA-N Glu-Asp-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(O)=O)NC(=O)[C@@H](N)CCC(O)=O JVSBYEDSSRZQGV-GUBZILKMSA-N 0.000 description 1
- ALCAUWPAMLVUDB-FXQIFTODSA-N Glu-Gln-Asn Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O ALCAUWPAMLVUDB-FXQIFTODSA-N 0.000 description 1
- MWMJCGBSIORNCD-AVGNSLFASA-N Glu-Leu-Leu Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O MWMJCGBSIORNCD-AVGNSLFASA-N 0.000 description 1
- MRWYPDWDZSLWJM-ACZMJKKPSA-N Glu-Ser-Asp Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(O)=O)C(O)=O MRWYPDWDZSLWJM-ACZMJKKPSA-N 0.000 description 1
- BYYNJRSNDARRBX-YFKPBYRVSA-N Gly-Gln-Gly Chemical compound NCC(=O)N[C@@H](CCC(N)=O)C(=O)NCC(O)=O BYYNJRSNDARRBX-YFKPBYRVSA-N 0.000 description 1
- AYBKPDHHVADEDA-YUMQZZPRSA-N Gly-His-Asn Chemical compound [H]NCC(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC(N)=O)C(O)=O AYBKPDHHVADEDA-YUMQZZPRSA-N 0.000 description 1
- AFWYPMDMDYCKMD-KBPBESRZSA-N Gly-Leu-Tyr Chemical compound NCC(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 AFWYPMDMDYCKMD-KBPBESRZSA-N 0.000 description 1
- JYGYNWYVKXENNE-OALUTQOASA-N Gly-Tyr-Trp Chemical compound [H]NCC(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(O)=O JYGYNWYVKXENNE-OALUTQOASA-N 0.000 description 1
- 102100021181 Golgi phosphoprotein 3 Human genes 0.000 description 1
- 241000219146 Gossypium Species 0.000 description 1
- 241000208818 Helianthus Species 0.000 description 1
- 235000003222 Helianthus annuus Nutrition 0.000 description 1
- VIVSWEBJUHXCDS-DCAQKATOSA-N His-Asn-Met Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CCSC)C(O)=O VIVSWEBJUHXCDS-DCAQKATOSA-N 0.000 description 1
- NNBWMLHQXBTIIT-HVTMNAMFSA-N His-Gln-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC1=CN=CN1)N NNBWMLHQXBTIIT-HVTMNAMFSA-N 0.000 description 1
- NELVFWFDOKRTOR-SDDRHHMPSA-N His-Gln-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC2=CN=CN2)N)C(=O)O NELVFWFDOKRTOR-SDDRHHMPSA-N 0.000 description 1
- HQKADFMLECZIQJ-HVTMNAMFSA-N His-Glu-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)O)NC(=O)[C@H](CC1=CN=CN1)N HQKADFMLECZIQJ-HVTMNAMFSA-N 0.000 description 1
- JJHWJUYYTWYXPL-PYJNHQTQSA-N His-Ile-Arg Chemical compound NC(N)=NCCC[C@@H](C(O)=O)NC(=O)[C@H]([C@@H](C)CC)NC(=O)[C@@H](N)CC1=CN=CN1 JJHWJUYYTWYXPL-PYJNHQTQSA-N 0.000 description 1
- CHIAUHSHDARFBD-ULQDDVLXSA-N His-Pro-Tyr Chemical compound C([C@H](N)C(=O)N1[C@@H](CCC1)C(=O)N[C@@H](CC=1C=CC(O)=CC=1)C(O)=O)C1=CN=CN1 CHIAUHSHDARFBD-ULQDDVLXSA-N 0.000 description 1
- 235000007340 Hordeum vulgare Nutrition 0.000 description 1
- 240000005979 Hordeum vulgare Species 0.000 description 1
- SCHZQZPYHBWYEQ-PEFMBERDSA-N Ile-Asn-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N SCHZQZPYHBWYEQ-PEFMBERDSA-N 0.000 description 1
- ADDYYRVQQZFIMW-MNXVOIDGSA-N Ile-Lys-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N ADDYYRVQQZFIMW-MNXVOIDGSA-N 0.000 description 1
- HXIDVIFHRYRXLZ-NAKRPEOUSA-N Ile-Ser-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(=O)O)N HXIDVIFHRYRXLZ-NAKRPEOUSA-N 0.000 description 1
- ONIBWKKTOPOVIA-BYPYZUCNSA-N L-Proline Chemical compound OC(=O)[C@@H]1CCCN1 ONIBWKKTOPOVIA-BYPYZUCNSA-N 0.000 description 1
- WHUUTDBJXJRKMK-VKHMYHEASA-N L-glutamic acid Chemical compound OC(=O)[C@@H](N)CCC(O)=O WHUUTDBJXJRKMK-VKHMYHEASA-N 0.000 description 1
- 101150022900 LFY gene Proteins 0.000 description 1
- 235000003228 Lactuca sativa Nutrition 0.000 description 1
- 240000008415 Lactuca sativa Species 0.000 description 1
- 101100254587 Lentinula edodes RPS1 gene Proteins 0.000 description 1
- RIMMMMYKGIBOSN-DCAQKATOSA-N Leu-Asn-Met Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CCSC)C(O)=O RIMMMMYKGIBOSN-DCAQKATOSA-N 0.000 description 1
- RRSLQOLASISYTB-CIUDSAMLSA-N Leu-Cys-Asp Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CS)C(=O)N[C@@H](CC(O)=O)C(O)=O RRSLQOLASISYTB-CIUDSAMLSA-N 0.000 description 1
- DLCXCECTCPKKCD-GUBZILKMSA-N Leu-Gln-Asn Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O DLCXCECTCPKKCD-GUBZILKMSA-N 0.000 description 1
- FQZPTCNSNPWHLJ-AVGNSLFASA-N Leu-Gln-Lys Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCCN)C(O)=O FQZPTCNSNPWHLJ-AVGNSLFASA-N 0.000 description 1
- KVMULWOHPPMHHE-DCAQKATOSA-N Leu-Glu-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O KVMULWOHPPMHHE-DCAQKATOSA-N 0.000 description 1
- QJUWBDPGGYVRHY-YUMQZZPRSA-N Leu-Gly-Cys Chemical compound CC(C)C[C@@H](C(=O)NCC(=O)N[C@@H](CS)C(=O)O)N QJUWBDPGGYVRHY-YUMQZZPRSA-N 0.000 description 1
- KGCLIYGPQXUNLO-IUCAKERBSA-N Leu-Gly-Glu Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCC(O)=O KGCLIYGPQXUNLO-IUCAKERBSA-N 0.000 description 1
- DBSLVQBXKVKDKJ-BJDJZHNGSA-N Leu-Ile-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C)C(O)=O DBSLVQBXKVKDKJ-BJDJZHNGSA-N 0.000 description 1
- QNBVTHNJGCOVFA-AVGNSLFASA-N Leu-Leu-Glu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CCC(O)=O QNBVTHNJGCOVFA-AVGNSLFASA-N 0.000 description 1
- LXKNSJLSGPNHSK-KKUMJFAQSA-N Leu-Leu-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)O)N LXKNSJLSGPNHSK-KKUMJFAQSA-N 0.000 description 1
- ZRHDPZAAWLXXIR-SRVKXCTJSA-N Leu-Lys-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O ZRHDPZAAWLXXIR-SRVKXCTJSA-N 0.000 description 1
- AIRUUHAOKGVJAD-JYJNAYRXSA-N Leu-Phe-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(O)=O)C(O)=O AIRUUHAOKGVJAD-JYJNAYRXSA-N 0.000 description 1
- DPURXCQCHSQPAN-AVGNSLFASA-N Leu-Pro-Pro Chemical compound CC(C)C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N1[C@H](C(O)=O)CCC1 DPURXCQCHSQPAN-AVGNSLFASA-N 0.000 description 1
- ADJWHHZETYAAAX-SRVKXCTJSA-N Leu-Ser-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N ADJWHHZETYAAAX-SRVKXCTJSA-N 0.000 description 1
- QWWPYKKLXWOITQ-VOAKCMCISA-N Leu-Thr-Leu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@H](C(O)=O)CC(C)C QWWPYKKLXWOITQ-VOAKCMCISA-N 0.000 description 1
- VHTIZYYHIUHMCA-JYJNAYRXSA-N Leu-Tyr-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O VHTIZYYHIUHMCA-JYJNAYRXSA-N 0.000 description 1
- YVMQJGWLHRWMDF-MNXVOIDGSA-N Lys-Gln-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CCCCN)N YVMQJGWLHRWMDF-MNXVOIDGSA-N 0.000 description 1
- NKKFVJRLCCUJNA-QWRGUYRKSA-N Lys-Gly-Lys Chemical compound NCCCC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCCN NKKFVJRLCCUJNA-QWRGUYRKSA-N 0.000 description 1
- WOEDRPCHKPSFDT-MXAVVETBSA-N Lys-His-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CC1=CN=CN1)NC(=O)[C@H](CCCCN)N WOEDRPCHKPSFDT-MXAVVETBSA-N 0.000 description 1
- QBEPTBMRQALPEV-MNXVOIDGSA-N Lys-Ile-Glu Chemical compound OC(=O)CC[C@@H](C(O)=O)NC(=O)[C@H]([C@@H](C)CC)NC(=O)[C@@H](N)CCCCN QBEPTBMRQALPEV-MNXVOIDGSA-N 0.000 description 1
- UQRZFMQQXXJTTF-AVGNSLFASA-N Lys-Lys-Glu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O UQRZFMQQXXJTTF-AVGNSLFASA-N 0.000 description 1
- MTBLFIQZECOEBY-IHRRRGAJSA-N Lys-Met-Lys Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCCCN)C(O)=O MTBLFIQZECOEBY-IHRRRGAJSA-N 0.000 description 1
- 241000616993 Magnoliidae Species 0.000 description 1
- 101710125418 Major capsid protein Proteins 0.000 description 1
- BLIPQDLSCFGUFA-GUBZILKMSA-N Met-Arg-Asn Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(N)=O)C(O)=O BLIPQDLSCFGUFA-GUBZILKMSA-N 0.000 description 1
- STTRPDDKDVKIDF-KKUMJFAQSA-N Met-Glu-Tyr Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 STTRPDDKDVKIDF-KKUMJFAQSA-N 0.000 description 1
- HAQLBBVZAGMESV-IHRRRGAJSA-N Met-Lys-Lys Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(O)=O HAQLBBVZAGMESV-IHRRRGAJSA-N 0.000 description 1
- XIGAHPDZLAYQOS-SRVKXCTJSA-N Met-Pro-Pro Chemical compound CSCC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N1[C@H](C(O)=O)CCC1 XIGAHPDZLAYQOS-SRVKXCTJSA-N 0.000 description 1
- XTSBLBXAUIBMLW-KKUMJFAQSA-N Met-Tyr-Glu Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N XTSBLBXAUIBMLW-KKUMJFAQSA-N 0.000 description 1
- 102000003792 Metallothionein Human genes 0.000 description 1
- 108090000157 Metallothionein Proteins 0.000 description 1
- 101100364971 Mus musculus Scai gene Proteins 0.000 description 1
- XZFYRXDAULDNFX-UHFFFAOYSA-N N-L-cysteinyl-L-phenylalanine Natural products SCC(N)C(=O)NC(C(O)=O)CC1=CC=CC=C1 XZFYRXDAULDNFX-UHFFFAOYSA-N 0.000 description 1
- PESQCPHRXOFIPX-UHFFFAOYSA-N N-L-methionyl-L-tyrosine Natural products CSCCC(N)C(=O)NC(C(O)=O)CC1=CC=C(O)C=C1 PESQCPHRXOFIPX-UHFFFAOYSA-N 0.000 description 1
- 229930193140 Neomycin Natural products 0.000 description 1
- 206010028980 Neoplasm Diseases 0.000 description 1
- 238000000636 Northern blotting Methods 0.000 description 1
- 101710141454 Nucleoprotein Proteins 0.000 description 1
- 108091034117 Oligonucleotide Proteins 0.000 description 1
- 108020005187 Oligonucleotide Probes Proteins 0.000 description 1
- 240000007594 Oryza sativa Species 0.000 description 1
- 235000007164 Oryza sativa Nutrition 0.000 description 1
- 206010034133 Pathogen resistance Diseases 0.000 description 1
- 244000115721 Pennisetum typhoides Species 0.000 description 1
- 235000007195 Pennisetum typhoides Nutrition 0.000 description 1
- BJEYSVHMGIJORT-NHCYSSNCSA-N Phe-Ala-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](C)NC(=O)[C@@H](N)CC1=CC=CC=C1 BJEYSVHMGIJORT-NHCYSSNCSA-N 0.000 description 1
- ILGCZYGFYQLSDZ-KKUMJFAQSA-N Phe-Ser-His Chemical compound N[C@@H](Cc1ccccc1)C(=O)N[C@@H](CO)C(=O)N[C@@H](Cc1cnc[nH]1)C(O)=O ILGCZYGFYQLSDZ-KKUMJFAQSA-N 0.000 description 1
- 241000758706 Piperaceae Species 0.000 description 1
- 235000010582 Pisum sativum Nutrition 0.000 description 1
- 240000004713 Pisum sativum Species 0.000 description 1
- 239000002202 Polyethylene glycol Substances 0.000 description 1
- FISHYTLIMUYTQY-GUBZILKMSA-N Pro-Gln-Gln Chemical compound NC(=O)CC[C@@H](C(O)=O)NC(=O)[C@H](CCC(N)=O)NC(=O)[C@@H]1CCCN1 FISHYTLIMUYTQY-GUBZILKMSA-N 0.000 description 1
- LXVLKXPFIDDHJG-CIUDSAMLSA-N Pro-Glu-Ser Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O LXVLKXPFIDDHJG-CIUDSAMLSA-N 0.000 description 1
- MCWHYUWXVNRXFV-RWMBFGLXSA-N Pro-Leu-Pro Chemical compound CC(C)C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@@H]2CCCN2 MCWHYUWXVNRXFV-RWMBFGLXSA-N 0.000 description 1
- XQPHBAKJJJZOBX-SRVKXCTJSA-N Pro-Lys-Glu Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O XQPHBAKJJJZOBX-SRVKXCTJSA-N 0.000 description 1
- HBBBLSVBQGZKOZ-GUBZILKMSA-N Pro-Met-Ala Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCSC)C(=O)N[C@@H](C)C(O)=O HBBBLSVBQGZKOZ-GUBZILKMSA-N 0.000 description 1
- WHNJMTHJGCEKGA-ULQDDVLXSA-N Pro-Phe-Leu Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(C)C)C(O)=O WHNJMTHJGCEKGA-ULQDDVLXSA-N 0.000 description 1
- OABLKWMLPUGEQK-JYJNAYRXSA-N Pro-Tyr-Met Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCSC)C(O)=O OABLKWMLPUGEQK-JYJNAYRXSA-N 0.000 description 1
- PGSWNLRYYONGPE-JYJNAYRXSA-N Pro-Val-Tyr Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O PGSWNLRYYONGPE-JYJNAYRXSA-N 0.000 description 1
- 101710083689 Probable capsid protein Proteins 0.000 description 1
- ONIBWKKTOPOVIA-UHFFFAOYSA-N Proline Natural products OC(=O)C1CCCN1 ONIBWKKTOPOVIA-UHFFFAOYSA-N 0.000 description 1
- 244000081426 Ranunculus ficaria Species 0.000 description 1
- 235000002226 Ranunculus ficaria Nutrition 0.000 description 1
- 244000088415 Raphanus sativus Species 0.000 description 1
- 235000006140 Raphanus sativus var sativus Nutrition 0.000 description 1
- 108020004511 Recombinant DNA Proteins 0.000 description 1
- 108700008625 Reporter Genes Proteins 0.000 description 1
- 108091028664 Ribonucleotide Proteins 0.000 description 1
- 108010003581 Ribulose-bisphosphate carboxylase Proteins 0.000 description 1
- 240000004808 Saccharomyces cerevisiae Species 0.000 description 1
- 235000007238 Secale cereale Nutrition 0.000 description 1
- 244000082988 Secale cereale Species 0.000 description 1
- OQPNSDWGAMFJNU-QWRGUYRKSA-N Ser-Gly-Tyr Chemical compound OC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 OQPNSDWGAMFJNU-QWRGUYRKSA-N 0.000 description 1
- CLKKNZQUQMZDGD-SRVKXCTJSA-N Ser-His-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)CO)CC1=CN=CN1 CLKKNZQUQMZDGD-SRVKXCTJSA-N 0.000 description 1
- LQESNKGTTNHZPZ-GHCJXIJMSA-N Ser-Ile-Asn Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(N)=O)C(O)=O LQESNKGTTNHZPZ-GHCJXIJMSA-N 0.000 description 1
- NNFMANHDYSVNIO-DCAQKATOSA-N Ser-Lys-Arg Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O NNFMANHDYSVNIO-DCAQKATOSA-N 0.000 description 1
- BYCVMHKULKRVPV-GUBZILKMSA-N Ser-Lys-Gln Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(N)=O)C(O)=O BYCVMHKULKRVPV-GUBZILKMSA-N 0.000 description 1
- NIOYDASGXWLHEZ-CIUDSAMLSA-N Ser-Met-Glu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCC(O)=O)C(O)=O NIOYDASGXWLHEZ-CIUDSAMLSA-N 0.000 description 1
- XNXRTQZTFVMJIJ-DCAQKATOSA-N Ser-Met-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O XNXRTQZTFVMJIJ-DCAQKATOSA-N 0.000 description 1
- NMZXJDSKEGFDLJ-DCAQKATOSA-N Ser-Pro-Lys Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CO)N)C(=O)N[C@@H](CCCCN)C(=O)O NMZXJDSKEGFDLJ-DCAQKATOSA-N 0.000 description 1
- QUGRFWPMPVIAPW-IHRRRGAJSA-N Ser-Pro-Phe Chemical compound OC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 QUGRFWPMPVIAPW-IHRRRGAJSA-N 0.000 description 1
- PIQRHJQWEPWFJG-UWJYBYFXSA-N Ser-Tyr-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](C)C(O)=O PIQRHJQWEPWFJG-UWJYBYFXSA-N 0.000 description 1
- 235000008515 Setaria glauca Nutrition 0.000 description 1
- FCHAMFUEENBIDH-UHFFFAOYSA-N Severin Natural products CC1CCC2C(C)C3CCC4(O)C(CC5C4CC(O)C6CC(CCC56C)OC(=O)C)C3CN2C1 FCHAMFUEENBIDH-UHFFFAOYSA-N 0.000 description 1
- 108020004682 Single-Stranded DNA Proteins 0.000 description 1
- 241000208255 Solanales Species 0.000 description 1
- 235000002597 Solanum melongena Nutrition 0.000 description 1
- 235000002595 Solanum tuberosum Nutrition 0.000 description 1
- 244000061456 Solanum tuberosum Species 0.000 description 1
- 235000011684 Sorghum saccharatum Nutrition 0.000 description 1
- 244000062793 Sorghum vulgare Species 0.000 description 1
- 235000009337 Spinacia oleracea Nutrition 0.000 description 1
- 244000300264 Spinacia oleracea Species 0.000 description 1
- 108091081024 Start codon Proteins 0.000 description 1
- 101001091268 Streptomyces hygroscopicus Hygromycin-B 7''-O-kinase Proteins 0.000 description 1
- 108700005078 Synthetic Genes Proteins 0.000 description 1
- BSNZTJXVDOINSR-JXUBOQSCSA-N Thr-Ala-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(O)=O BSNZTJXVDOINSR-JXUBOQSCSA-N 0.000 description 1
- VBPDMBAFBRDZSK-HOUAVDHOSA-N Thr-Asn-Trp Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)O)N)O VBPDMBAFBRDZSK-HOUAVDHOSA-N 0.000 description 1
- VTVVYQOXJCZVEB-WDCWCFNPSA-N Thr-Leu-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O VTVVYQOXJCZVEB-WDCWCFNPSA-N 0.000 description 1
- 108010022394 Threonine synthase Proteins 0.000 description 1
- 102000006601 Thymidine Kinase Human genes 0.000 description 1
- 108020004440 Thymidine kinase Proteins 0.000 description 1
- 108700009124 Transcription Initiation Site Proteins 0.000 description 1
- 241000219793 Trifolium Species 0.000 description 1
- 235000021307 Triticum Nutrition 0.000 description 1
- 244000098338 Triticum aestivum Species 0.000 description 1
- PXQPYPMSLBQHJJ-WFBYXXMGSA-N Trp-Asp-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H](CC1=CNC2=CC=CC=C21)N PXQPYPMSLBQHJJ-WFBYXXMGSA-N 0.000 description 1
- DWJQKEZKLQCHKO-SRVKXCTJSA-N Tyr-Asn-Cys Chemical compound C1=CC(=CC=C1C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CS)C(=O)O)N)O DWJQKEZKLQCHKO-SRVKXCTJSA-N 0.000 description 1
- XQYHLZNPOTXRMQ-KKUMJFAQSA-N Tyr-Glu-Arg Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O XQYHLZNPOTXRMQ-KKUMJFAQSA-N 0.000 description 1
- ZRPLVTZTKPPSBT-AVGNSLFASA-N Tyr-Glu-Ser Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O ZRPLVTZTKPPSBT-AVGNSLFASA-N 0.000 description 1
- DWAMXBFJNZIHMC-KBPBESRZSA-N Tyr-Leu-Gly Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O DWAMXBFJNZIHMC-KBPBESRZSA-N 0.000 description 1
- LUMQYLVYUIRHHU-YJRXYDGGSA-N Tyr-Ser-Thr Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(O)=O LUMQYLVYUIRHHU-YJRXYDGGSA-N 0.000 description 1
- MDXLPNRXCFOBTL-BZSNNMDCSA-N Tyr-Ser-Tyr Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O MDXLPNRXCFOBTL-BZSNNMDCSA-N 0.000 description 1
- PVPAOIGJYHVWBT-KKHAAJSZSA-N Val-Asn-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](C(C)C)N)O PVPAOIGJYHVWBT-KKHAAJSZSA-N 0.000 description 1
- PDDJTOSAVNRJRH-UNQGMJICSA-N Val-Thr-Phe Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H](C(C)C)N)O PDDJTOSAVNRJRH-UNQGMJICSA-N 0.000 description 1
- 235000002096 Vicia faba var. equina Nutrition 0.000 description 1
- 108020005202 Viral DNA Proteins 0.000 description 1
- 241000700605 Viruses Species 0.000 description 1
- 108010027570 Xanthine phosphoribosyltransferase Proteins 0.000 description 1
- 235000005824 Zea mays ssp. parviglumis Nutrition 0.000 description 1
- 235000016383 Zea mays subsp huehuetenangensis Nutrition 0.000 description 1
- 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 1
- 230000001594 aberrant effect Effects 0.000 description 1
- 239000002253 acid Substances 0.000 description 1
- 150000007513 acids Chemical class 0.000 description 1
- 230000009471 action Effects 0.000 description 1
- 239000000654 additive Substances 0.000 description 1
- 230000000996 additive effect Effects 0.000 description 1
- 108010087924 alanylproline Proteins 0.000 description 1
- 230000004075 alteration Effects 0.000 description 1
- 229940126575 aminoglycoside Drugs 0.000 description 1
- 239000003242 anti bacterial agent Substances 0.000 description 1
- 238000013459 approach Methods 0.000 description 1
- 108010093581 aspartyl-proline Proteins 0.000 description 1
- 230000003416 augmentation Effects 0.000 description 1
- 239000002363 auxin Substances 0.000 description 1
- 239000011324 bead Substances 0.000 description 1
- 150000008331 benzenesulfonamides Chemical class 0.000 description 1
- 239000001511 capsicum annuum Substances 0.000 description 1
- 210000002421 cell wall Anatomy 0.000 description 1
- 235000013339 cereals Nutrition 0.000 description 1
- 239000003795 chemical substances by application Substances 0.000 description 1
- 239000013599 cloning vector Substances 0.000 description 1
- 229910001431 copper ion Inorganic materials 0.000 description 1
- 235000005822 corn Nutrition 0.000 description 1
- 238000005520 cutting process Methods 0.000 description 1
- 101150064332 cyc07 gene Proteins 0.000 description 1
- 239000004062 cytokinin Substances 0.000 description 1
- UQHKFADEQIVWID-UHFFFAOYSA-N cytokinin Natural products C1=NC=2C(NCC=C(CO)C)=NC=NC=2N1C1CC(O)C(CO)O1 UQHKFADEQIVWID-UHFFFAOYSA-N 0.000 description 1
- 230000007547 defect Effects 0.000 description 1
- 239000005547 deoxyribonucleotide Substances 0.000 description 1
- 125000002637 deoxyribonucleotide group Chemical group 0.000 description 1
- 238000001514 detection method Methods 0.000 description 1
- 230000004069 differentiation Effects 0.000 description 1
- 102000004419 dihydrofolate reductase Human genes 0.000 description 1
- 235000005489 dwarf bean Nutrition 0.000 description 1
- 244000013123 dwarf bean Species 0.000 description 1
- 230000005014 ectopic expression Effects 0.000 description 1
- 238000004520 electroporation Methods 0.000 description 1
- 210000002257 embryonic structure Anatomy 0.000 description 1
- 239000003623 enhancer Substances 0.000 description 1
- 238000002474 experimental method Methods 0.000 description 1
- 239000013604 expression vector Substances 0.000 description 1
- 239000003337 fertilizer Substances 0.000 description 1
- 230000004883 flower formation Effects 0.000 description 1
- BRZYSWJRSDMWLG-CAXSIQPQSA-N geneticin Natural products O1C[C@@](O)(C)[C@H](NC)[C@@H](O)[C@H]1O[C@@H]1[C@@H](O)[C@H](O[C@@H]2[C@@H]([C@@H](O)[C@H](O)[C@@H](C(C)O)O2)N)[C@@H](N)C[C@H]1N BRZYSWJRSDMWLG-CAXSIQPQSA-N 0.000 description 1
- 210000004602 germ cell Anatomy 0.000 description 1
- 239000003862 glucocorticoid Substances 0.000 description 1
- 235000013922 glutamic acid Nutrition 0.000 description 1
- 239000004220 glutamic acid Substances 0.000 description 1
- 108010078144 glutaminyl-glycine Proteins 0.000 description 1
- 108010050475 glycyl-leucyl-tyrosine Proteins 0.000 description 1
- 239000001963 growth medium Substances 0.000 description 1
- 230000036541 health Effects 0.000 description 1
- 230000002363 herbicidal effect Effects 0.000 description 1
- 239000004009 herbicide Substances 0.000 description 1
- 239000008241 heterogeneous mixture Substances 0.000 description 1
- 230000006872 improvement Effects 0.000 description 1
- 238000011065 in-situ storage Methods 0.000 description 1
- 230000000415 inactivating effect Effects 0.000 description 1
- 230000002779 inactivation Effects 0.000 description 1
- SEOVTRFCIGRIMH-UHFFFAOYSA-N indole-3-acetic acid Chemical compound C1=CC=C2C(CC(=O)O)=CNC2=C1 SEOVTRFCIGRIMH-UHFFFAOYSA-N 0.000 description 1
- 230000010354 integration Effects 0.000 description 1
- 238000002955 isolation Methods 0.000 description 1
- 108010034529 leucyl-lysine Proteins 0.000 description 1
- 108010091871 leucylmethionine Proteins 0.000 description 1
- 235000009973 maize Nutrition 0.000 description 1
- 210000001161 mammalian embryo Anatomy 0.000 description 1
- 108010083942 mannopine synthase Proteins 0.000 description 1
- 238000013507 mapping Methods 0.000 description 1
- 239000011159 matrix material Substances 0.000 description 1
- 230000007246 mechanism Effects 0.000 description 1
- 239000012528 membrane Substances 0.000 description 1
- 230000000442 meristematic effect Effects 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
- 108010056582 methionylglutamic acid Proteins 0.000 description 1
- 108010005942 methionylglycine Proteins 0.000 description 1
- 238000000520 microinjection Methods 0.000 description 1
- 239000000203 mixture Substances 0.000 description 1
- 238000010369 molecular cloning Methods 0.000 description 1
- 238000012544 monitoring process Methods 0.000 description 1
- 230000000877 morphologic effect Effects 0.000 description 1
- 239000006870 ms-medium Substances 0.000 description 1
- 230000010115 negative regulation of flower development Effects 0.000 description 1
- 229960004927 neomycin Drugs 0.000 description 1
- 230000009871 nonspecific binding Effects 0.000 description 1
- 239000002751 oligonucleotide probe Substances 0.000 description 1
- 239000012188 paraffin wax Substances 0.000 description 1
- 230000035515 penetration Effects 0.000 description 1
- 108010051242 phenylalanylserine Proteins 0.000 description 1
- 230000035479 physiological effects, processes and functions Effects 0.000 description 1
- 230000008121 plant development Effects 0.000 description 1
- 230000037039 plant physiology Effects 0.000 description 1
- 229920001223 polyethylene glycol Polymers 0.000 description 1
- 229920000642 polymer Polymers 0.000 description 1
- 235000012015 potatoes Nutrition 0.000 description 1
- 238000001556 precipitation Methods 0.000 description 1
- 239000002243 precursor Substances 0.000 description 1
- 238000007639 printing Methods 0.000 description 1
- 230000035755 proliferation Effects 0.000 description 1
- 108010031719 prolyl-serine Proteins 0.000 description 1
- 108010090894 prolylleucine Proteins 0.000 description 1
- 230000009467 reduction Effects 0.000 description 1
- 238000011160 research Methods 0.000 description 1
- 230000004044 response Effects 0.000 description 1
- 238000010839 reverse transcription Methods 0.000 description 1
- 239000002336 ribonucleotide Substances 0.000 description 1
- 125000002652 ribonucleotide group Chemical group 0.000 description 1
- 235000009566 rice Nutrition 0.000 description 1
- 230000005070 ripening Effects 0.000 description 1
- 108010026333 seryl-proline Proteins 0.000 description 1
- 230000035939 shock Effects 0.000 description 1
- 235000020354 squash Nutrition 0.000 description 1
- 210000000130 stem cell Anatomy 0.000 description 1
- 230000001629 suppression Effects 0.000 description 1
- 229940037128 systemic glucocorticoids Drugs 0.000 description 1
- 238000012360 testing method Methods 0.000 description 1
- 238000012090 tissue culture technique Methods 0.000 description 1
- 238000013518 transcription Methods 0.000 description 1
- 230000035897 transcription Effects 0.000 description 1
- 230000002103 transcriptional effect Effects 0.000 description 1
- 230000001131 transforming effect Effects 0.000 description 1
- 230000007704 transition Effects 0.000 description 1
- 238000011144 upstream manufacturing Methods 0.000 description 1
- 235000013311 vegetables Nutrition 0.000 description 1
- 230000008511 vegetative development Effects 0.000 description 1
- 230000001018 virulence Effects 0.000 description 1
- 230000000007 visual effect Effects 0.000 description 1
- 238000012800 visualization Methods 0.000 description 1
Images
Classifications
-
- 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
- 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
-
- 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
- C12N15/8262—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield involving plant development
- C12N15/827—Flower development or morphology, e.g. flowering promoting factor [FPF]
-
- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
- Y02A40/146—Genetically Modified [GMO] plants, e.g. transgenic plants
Definitions
- the invention relates generally to plant genetic engineering, and specifically to novel genetically engineered plants characterized as having a phenotype of early flower meristem development, and methods for producing such plants.
- angiosperm species are induced to flower in response to environmental stimuli such as day length and temperature, and internal cues, such as age.
- Adult organs of flowering plants develop from groups of stem cells called meristems.
- the identity of a meristem is inferred from structures it produces: vegetative meristems give rise to roots and leaves, inflorescence meristems give rise to flower meristems, and flower meristems give rise to floral organs such as sepals and petals.
- vegetative meristems give rise to roots and leaves
- inflorescence meristems give rise to flower meristems
- flower meristems give rise to floral organs such as sepals and petals.
- meristems capable of generating new meristems of different identity, but their own identity can change during development.
- the homologue of the Arabidopsis LEAFY gene is FLORICAULA (Coen, et al., Cell, 63:1311, 1990) and that of the APETALA1 gene is SQUAMOSA (Huijser, et al., EMBO J, 11:1239, 1992).
- FLORICAULA Coen, et al., Cell, 63:1311, 1990
- SQUAMOSA Huijser, et al., EMBO J, 11:1239, 1992.
- the latter pair contains MADS box domains.
- LEAFY is expressed very early in floral anlagen and floral primordia, consistent with it having a direct role in establishing floral meristem identity. In the developing floral primordium, LEAFY expression is detected much earlier than expression of the homeotic genes AG and AP3, suggesting that LEAFY plays a role in controlling the expression of floral homeotic genes.
- the present invention arose out of the discovery that a genetically modified plant cell could be produced, from which a whole plant can be regenerated which stably incorporates a flower development genetic trait introduced into the plant cell. Specifically, the trait of early flowering can be imparted on a plant by genetic modification according to the method of the invention.
- a method of producing a genetically modified plant characterized as having modulated flower meristem development comprises contacting a plant cell with a vector(s), comprising a nucleic acid sequence comprising at least one structural gene encoding a protein for modulating flower meristem development, operably associated with a promoter to obtain a transformed plant cell; producing plants from said transformed plant cell; and selecting a plant exhibiting modulated flower meristem development.
- FIG. 1 shows a schematic illustration of pDW139, which is the parental plasmid for construction of 35S::LFY vectors. Open reading frame of LEAFY(LFY) is hatched; 5′ and 3′ untranslated regions are stippled.
- FIG. 2 shows the early flowering phenotype of 35S::LFY tobacco plants. Left, control plant, transformed with an unrelated construct. Middle and right, two independently derived T 2 plants carrying a 35S::LFY transgene (lines 146.21, 146.26). Plants are five weeks old.
- FIG. 3 shows precocious enlargement of apical meristem in 35S::LFY tobacco plants.
- Panel (A) Control, transformed with an unrelated construct.
- Panel (B) Experimental plant, transformed with a 35S::LFY construct. Size bar, 50 ⁇ m.
- FIG. 4 shows the early flowering phenotype of 35S::LFY Arabidopsis plants.
- Panel (A) Control plant, transformed with an unrelated construct.
- the rosette leaves (rl) are significantly larger than the cotyledons (cot).
- Panel (B) 35S::LFY transformant (line 151.106).
- the first two rosette leaves (rl) are smaller than the cotyledons.
- FIG. 5 shows the conversion of all shoots into flowers in 35S::LFY Arabidopsis plants.
- Panel (A) For comparison, a drawing of a mature Arabidopsis plant (Nossen ecotype) of about six weeks of age is shown.
- Panel (B) Top view of a wild-type Arabidopsis inflorescence, illustrating the indeterminacy of the shoot meristem.
- Panels (C)-(E) show 35S::LFY plants (generated in the Nossen ecotype), three weeks old.
- Panel (C) Replacement of shoots with single flowers (triangles) (line 151.201). A cotyledon is indicated (cot).
- FIG. 6 shows constitutive expression of Arabidopsis LFY converts aspen shoots into flowers.
- Panels a and b show five-month-old shoots of hybrid aspen ( Populous tremula x tremuloides ) grown in tissue culture.
- Panel a shows a 35S::LFY transformant. Solitary, lateral flowers in the axils of leaves (lf) and an abnormal terminal flower (tf) are indicated.
- Panel b shows a non-transgenic control. Arrowheads indicate axils of leaves, from which lateral vegetative shoots will emerge, normally in the following year.
- FIG. 7 shows 35S::LFY phenotype is partly suppressed by an ap1 mutation.
- Panel a shows five-week-old plants that carry the erecta mutation.
- the 35S::LFY AP1 + plant (left) has no elongated primary shoot.
- a primary shoot is well developed in the 35S::LFY ap1 plant (middle), although the primary shoot still terminates prematurely, and is shorter than that of the non-transgenic ap1 plant (right).
- Panels b-d show a detailed view of 35S::LFY ap1 plants.
- Panel b shows a close-up view of lateral shoot indicated by arrowhead in panel a.
- Examples of monocotyledonous plants include, but are not limited to, asparagus, field and sweet corn, barley, wheat, rice, sorghum, onion, pearl millet, rye and oats.
- Examples of dicotyledonous plants include, but are not limited to tomato, tobacco, cotton, rapeseed, field beans, soybeans, peppers, lettuce, peas, alfalfa, clover, cole crops or Brassica oleracea (e.g., cabbage, broccoli, cauliflower, brussel sprouts), radish, carrot, beets, eggplant, spinach, cucumber, squash, melons, cantaloupe, sunflowers and various ornamentals.
- Exemplary models described herein include the tobacco plant and the perennial tree, aspen.
- heterologous nucleic acid sequence refers to at least one structural gene operably associated with a regulatory sequence such as a promoter.
- the nucleic acid sequence originates in a foreign species, or, in the same species if substantially modified from its original form.
- heterologous nucleic acid sequence includes a nucleic acid originating in the same species, where such sequence is operably linked to a promoter that differs from the natural or wild-type promoter.
- nucleic acid sequence refers to a polymer of deoxyribonucleotides or ribonucleotides, in the form of a separate fragment or as a component of a larger construct.
- DNA encoding the proteins utilized in the method of the invention can be assembled from cDNA fragments or from oligonucleotides which provide a synthetic gene which is capable of being expressed in a recombinant transcriptional unit.
- Polynucleotide or nucleic acid sequences of the invention include DNA, RNA and cDNA sequences.
- Examples of structural genes that may be employed in the present invention include the LEAFY gene and the APETALA1 gene which control flowering. Also included in the present invention are structural and functional homologues of the LEAFY and APETALA1 genes. For example, in Antirrhinum majus, the snapdragon, the homologue of the LEAFY gene is the FLORICAULA gene and the homologue of the APETALA1 gene is the SQUAMOSA gene. Other genes which control flowering will be known to those of skill in the art or can be readily ascertained.
- Nucleic acid sequences utilized in the invention can be obtained by several methods.
- the DNA can be isolated using hybridization procedures which are well known in the art. These include, but are not limited to: 1) hybridization of probes to genomic or cDNA libraries to detect shared nucleotide sequences; 2) antibody screening of expression libraries to detect shared structural features and 3) synthesis by the polymerase chain reaction (PCR). Sequences for specific genes can also be found in GenBank, National Institutes of Health computer database.
- Hybridization procedures useful for screening for desired nucleic acid sequences utilized herein employ labeled mixed synthetic oligonucleotide probes where each probe is potentially the complete complement of a specific DNA sequence in the hybridization sample which includes a heterogeneous mixture of denatured double-stranded DNA.
- hybridization is preferably performed on either single-stranded DNA or denatured double-stranded DNA.
- Hybridization is particularly useful in the detection of cDNA clones derived from sources where an extremely low amount of mRNA sequences relating to the polypeptide of interest are present.
- a cDNA expression library such as lambda gtl 11, can be screened indirectly for a heterologous polypeptide having at least one epitope, using antibodies specific for the heterologous protein.
- antibodies can be either polyclonally or monoclonally derived and used to detect expression product indicative of the presence of heterologous protein cDNA.
- a polypeptide sequence can be deduced from the genetic code, however, the degeneracy of the code must be taken into account.
- Nucleic acid sequences utilized in the invention include sequences which are degenerate as a result of the genetic code. There are 20 natural amino acids, most of which are specified by more than one codon. Therefore, as long as the amino acid sequence of heterologous polypeptide results in a functional polypeptide (at least, in the case of the sense polynucleotide strand), all degenerate nucleotide sequences are included in the invention.
- the heterologous nucleic acid sequences utilized herein are structural genes for flower meristem development.
- such genes encode a protein that is sufficient for the initiation of flowering, and most preferably, the nucleic acid sequence encodes the LEAFY protein.
- the LEAFY gene or other flower meristem development gene may be utilized alone or in combination with another structural gene, such as another gene which encodes a protein important in the development of flowering.
- An example of such a gene is the APETALA1 gene.
- Genetically modified plants of the present are produced by contacting a plant cell with a vector comprising a heterologous nucleic acid sequence comprising at least one structural gene encoding a protein that modulates flower meristem development.
- the structural gene of interest must be operably associated with a promoter which is effective in the plant cells to cause transcription of the gene of interest.
- a polyadenylation sequence or transcription control sequence also recognized in plant cells may also be employed. It is preferred that the vector harboring the heterologous nucleic acid sequence also contain one or more selectable marker genes so that the transformed cells can be selected from non-transformed cells in culture, as described herein.
- operably associated refers to functional linkage between a promoter sequence and the structural gene regulated by the promoter nucleic acid sequence.
- the operably linked promoter controls the expression of the polypeptide encoded by the structural gene.
- the expression of structural genes employed in the present invention may be driven by a number of promoters.
- the endogenous promoter of a structural gene of interest may be utilized for transcriptional regulation of the gene, preferably, the promoter is a foreign regulatory sequence.
- suitable viral promoters include the 35S RNA and 19S RNA promoters of CaMV (Brisson, et al., Nature, 310:511, 1984; Odell, et al., Nature, 313:810, 1985); the full-length transcript promoter from Figwort Mosaic Virs (FMV) (Gowda, et al., J Cell Biochem., 13D: 301, 1989) and the coat protein promoter to TMV (Takamatsu, et al., EMBO J. 3:17, 1987).
- CaMV 35S RNA and 19S RNA promoters of CaMV
- FMV Figwort Mosaic Virs
- TMV coat protein promoter to TMV
- Promoters useful in the invention include both constitutive and inducible natural promoters as well as engineered promoters.
- the CaMV promoters are examples of constitutive promoters.
- an inducible promoter should 1) provide low expression in the absence of the inducer; 2) provide high expression in the presence of the inducer; 3) use an induction scheme that does not interfere with the normal physiology of the plant; and 4) have no effect on the expression of other genes.
- inducible promoters useful in plants include those induced by chemical means, such as the yeast metallothionein promoter which is activated by copper ions (Mett, et al., Proc. Nat. Acad.
- the particular promoter selected should be capable of causing sufficient expression to result in the production of an effective amount of the structural gene product, e.g., LEAFY, to cause early floral meristem development:
- the promoters used in the vector constructs of the present invention may be modified, if desired, to affect their control characteristics.
- a selectable marker may be associated with the heterologous nucleic acid sequence, i.e., the structural gene operably linked to a promoter.
- the term “marker” refers to a gene encoding a trait or a phenotype which permits the selection of, or the screening for, a plant or plant cell containing the marker.
- the marker gene is an antibiotic resistance gene whereby the appropriate antibiotic can be used to select for transformed cells from among cells that are not transformed.
- Suitable selectable markers include adenosine deaminase, dihydrofolate reductase, hygromycin-B-phosphotransferase, thymidine kinase, xanthine-guanine phospho-ribosyltransferase and amino-glycoside 3′-O-phosphotransferase II (kanamycin, neomycin and G418 resistance).
- Other suitable markers will be known to those of skill in the art.
- Vector(s) employed in the present invention for transformation of a plant cell to modulate flower meristem development comprise a nucleic acid sequence comprising at least one structural gene encoding a protein that modulates flower meristem development, operably associated with a promoter.
- a suitable vector To commence a transformation process in accordance with the present invention, it is first necessary to construct a suitable vector and properly introduce it into the plant cell. The details of the construction of the vectors then utilized herein are known to those skilled in the art of plant genetic engineering.
- the heterologous nucleic acid sequences utilized in the present invention can be introduced into plant cells using Ti plasmids, root-inducing (Ri) plasmids, and plant virus vectors.
- Ti plasmids root-inducing (Ri) plasmids
- plant virus vectors for reviews of such techniques see, for example, Weissbach & Weissbach, 1988, Methods for Plant Molecular Biology, Academic Press, NY, Section VIII, pp. 421-463; and Grierson & Corey, 1988, Plant Molecular Biology, 2d Ed., Blackie, London, Ch. 7-9, and Horsch, et al., Science, 227:1229, 1985, both incorporated herein by reference).
- transformation of plants in accordance with the invention may be carried out in essentially any of the various ways known to those skilled in the art of plant molecular biology. (See, for example, Methods of Enzymology, Vol. 153, 1987, Wu and Grossman, Eds., Academic Press, incorporated herein by reference).
- transformation means alteration of the genotype of a host plant by the introduction of a heterologous nucleic acid sequence.
- a heterologous nucleic acid sequence can be introduced into a plant cell utilizing Agrobacterium tumefaciens containing the Ti plasmid.
- Agrobacterium tumefaciens culture it is most advantageous to use a non-oncogenic strain of the Agrobacterium as the vector carrier so that normal non-oncogenic differentiation of the transformed tissues is possible.
- the Agrobacterium harbor a binary Ti plasmid system.
- Such a binary system comprises 1) a first Ti plasmid having a virulence region essential for the introduction of transfer DNA (T-DNA) into plants, and 2) a chimeric plasmid.
- the latter contains at least one border region of the T-DNA region of a wild-type Ti plasmid flanking the nucleic acid to be transferred.
- Binary Ti plasmid systems have been shown effective to transform plant cells (De Framond, Biotechnology, 1:262, 1983; Hoekema, et al., Nature, 303:179, 1983). Such a binary system is preferred because it does not require integration into Ti plasmid in Agrobacterium.
- Methods involving the use of Agrobacterium include, but are not limited to: 1) co-cultivation of Agrobacterium with cultured isolated protoplasts; 2) transformation of plant cells or tissues with Agrobacterium; or 3) transformation of seeds, apices or meristems with Agrobacterium.
- gene transfer can be accomplished by in situ transformation by Agrobacterium, as described by Bechtold, et al., ( C.R. Acad. Sci. Paris, 316:1194, 1993) and exemplified in the Examples herein. This approach is based on the vacuum infiltration of a suspension of Agrobacterium cells.
- a preferred method of introducing heterologous nucleic acid into plant cells is to infect such plant cells, an explant, a meristem or a seed, with transformed Agrobacterium tumefaciens as described above. Under appropriate conditions known in the art, the transformed plant cells are grown to form shoots, roots, and develop further into plants.
- a preferred vector(s) of the invention comprises a Ti plasmid binary system wherein the heterologous nucleic acid sequence encodes the LEAFY protein.
- Such a vector may optionally contain a nucleic acid sequence which encodes a second flower development factor, such as APETALA1.
- two vectors can be utilized wherein each vector contains a heterologous nucleic acid sequence.
- Other flower development genes can be utilized for construction of one or more vectors, in a similar manner.
- heterologous nucleic acid can be introduced into a plant cell by contacting the plant cell using mechanical or chemical means.
- the nucleic acid can be mechanically transferred by microinjection directly into plant cells by use of micropipettes.
- the nucleic acid may be transferred into the plant cell by using polyethylene glycol which forms a precipitation complex with genetic material that is taken up by the cell.
- nucleic acid Another method for introducing nucleic acid into a plant cell is high velocity ballistic penetration by small particles with the nucleic acid to be introduced contained either within the matrix of small beads or particles, or on the surface thereof (Klein, et al., Nature 327:70, 1987). Although, typically only a single introduction of a new nucleic acid sequence is required, this method particularly provides for multiple introductions.
- Cauliflower mosaic virus may also be used as a vector for introducing heterologous nucleic acid into plant cells (U.S. Pat. No. 4,407,956).
- CaMV viral DNA genome is inserted into a parent bacterial plasmid creating a recombinant DNA molecule which can be propagated in bacteria.
- the recombinant plasmid again may be cloned and further modified by introduction of the desired nucleic acid sequence.
- the modified viral portion of the recombinant plasmid is then excised from the parent bacterial plasmid, and used to inoculate the plant cells or plants.
- contacting refers to any means of introducing the vector(s) into the plant cell, including chemical and physical means as described above.
- contacting refers to introducing the nucleic acid or vector into plant cells (including an explant, a meristem or a seed), via Agrobacterium tumefaciens transformed with the heterologous nucleic acid as described above.
- Regeneration from protoplasts varies from species to species of plants, but generally a suspension of protoplasts is first made. In certain species, embryo formation can then be induced from the protoplast suspension, to the stage of ripening and germination as natural embryos.
- the culture media will generally contain various amino acids and hormones, necessary for growth and regeneration. Examples of hormones utilized include auxin and cytokinins. It is sometimes advantageous to add glutamic acid and proline to the medium, especially for such species as corn and alfalfa. Efficient regeneration will depend on the medium, on the genotype, and on the history of the culture. If these variables are controlled, regeneration is reproducible.
- the mature transgenic plants are propagated by the taking of cuttings or by tissue culture techniques to produce multiple identical plants. Selection of desirable transgenotes is made and new varieties are obtained and propagated vegetatively for commercial use.
- the mature transgenic plants can be self crossed to produce a homozygous inbred plant. The inbred plant produces seed containing the newly introduced foreign gene(s). These seeds can be grown to produce plants that would produce the selected phenotype, e.g. early flowering.
- Parts obtained from the regenerated plant such as flowers, seeds, leaves, branches, fruit, and the like are included in the invention, provided that these parts comprise cells that have been transformed as described. Progeny and variants, and mutants of the regenerated plants are also included within the scope of the invention, provided that these parts comprise the introduced nucleic acid sequences.
- the LEAFY protein is likely to be a transcription factor, as it localizes to the nucleus and can bind to DNA in vitro.
- most other floral regulatory genes including APETALA1, encode known transcription factors with a MADS DNA-binding domain (e.g., Mandel, et al., Nature, 360:273, 1992).
- LFY ectopic LEAFY
- CaMV cauliflower mosaic virus
- the chimeric 35S::LFY gene was introduced into Arabidopsis and the distantly related tobacco plants by T-DNA mediated transformation.
- Other examples show aspen trees transformed with LFY.
- the phenotypic effects observed in transgenic plants show that LFY is not only necessary, but also sufficient for the initiation of flower development.
- the pDW146 vector was used for transformation of tobacco.
- the pDW151 vector was used for transformation of Arabidopsis. Both vectors are derived from plasmid pDW139, which contains the entire open reading frame of the LEAFY (LFY) gene from Arabidopsis thaliana (Weigel, D., et al., Cell, 69:843, 1992), plus 21 bp upstream of the initiation codon and 195 bp downstream of the stop codon (for cDNA sequence, see Weigel et al., supra; genomic sequence deposited in GenBank under accession number M91208).
- LEAFY LEAFY
- a Bg12 site was added by polymerase chain reaction (Saiki, et al., Science, 239:487, 1988).
- a genomic ScaI site was eliminated in the cloning process, and it is followed immediately by an Asp718 site derived from the pBluescript KS+cloning vector (FIG. 1).
- FIG. 1 shows a schematic illustration of pDW139 parental plasmid for construction of 35S::LFY vectors.
- LEAFY LEAFY
- pDW146 To construct pDW146, the 1.5 kb Bg12/Asp718 fragment carrying the LFY sequences was inserted into the binary T-DNA transformation vector pMON530 (Rogers, et al., Meth. Enzymol, 153:253, 1987), using the same sites in the vector.
- This vector contains an expression cassette comprising a 0.3 kb fragment of the cauliflower mosaic virus 35S promoter, including the transcription initiation site (Guilley, et al., Cell, 30:763, 1982); Odell, et al., supra.); a multilinker containing several unique restriction sites; and a functional polyadenylation signal from the Ti plasmid T-DNA nopaline synthase gene (“3′ nos”; [Bevan, et al., Plant Cell, 1:141, 1983]).
- pDW151 To construct pDW151, the Asp718 site of pDW139 was filled in with Klenow enzyme (Sambrook, et al., Molecular Cloning 2nd ed. (Cold Spring Harbor: Cold Spring Harbor Laboratory, 1989) and a Bg12 linker was added. The resulting Bg12 fragment was inserted into the BamH1 site of pCGN18, a transformation vector containing a CaMV 35S promoter 3′ nos expression cassette (Jack, et al, Cell, 76:703, 1994).
- Klenow enzyme Standardbrook, et al., Molecular Cloning 2nd ed. (Cold Spring Harbor: Cold Spring Harbor Laboratory, 1989) and a Bg12 linker was added.
- the resulting Bg12 fragment was inserted into the BamH1 site of pCGN18, a transformation vector containing a CaMV 35S promoter 3′ nos expression cassette (Jack, et al, Cell, 76:703, 1994).
- pDW146 and pDW151 plasmid DNAs isolated from E. coli were transformed into Agrobacterium tumefaciens strain LBA4404 (Ooms, et al, Plasmid, 7:15, 1982) or ASE (Fraley, et al, Biotechnology 3:629, 1985), respectively, using the freeze-thaw method as described (Hofgen and Willmitzer, Nucl. Acids Res, 16:9877, 1988), except that LB medium (Sambrook, et al., supra) was used instead of YEB.
- FIG. 2 shows the early flowering phenotype of 35S::LFY tobacco plants.
- the left panel shows a control plant, transformed with an unrelated construct (a LFY promoter fused to a GUS reporter gene).
- the middle and right panels show two independently derived T 2 plants carrying a 35S::LFY transgene (lines 146.21, 146.26).
- the plants shown are five weeks old. Note abundant proliferation of leaves in the control, while the experimental plants have produced only two true leaves before initiating a terminal flower.
- the insert shows a top view of floral bud of plant shown at the right. The bud is still unopened.
- transgenic plants exhibit the same dramatic phenotype in the progeny of the primary transformants (T 2 generation).
- Transgenic plants develop only one pair of true leaves, in addition to the embryonic leaves (cotyledons), before they produce a terminal flower (FIG. 2).
- wild-type tobacco plants also produce a terminal flower, they generate twenty to twenty-five pairs of leaves before flowering.
- constitutive LFY expression causes precocious conversion of the shoot meristem into a floral meristem.
- Histological sections of transgenic plants reveal that the apical meristem is morphologically different from that of untransformed plants at least as early as five days after germination (FIG. 3). The result of these changes is that transformed plants produce visible floral buds after two weeks, while normal tobacco plants flower only after about three to five months (the exact time depends on environmental conditions, such as light intensity, fertilizer, size of pots in which plants are grown, etc.).
- FIG. 3 shows precocious enlargement of apical meristem in 35S::LFY tobacco plants.
- Panel (A) is a control, transformed with the unrelated construct described in FIG. 2.
- Panel (B) shows an experimental plant, transformed with a 35S::LFY construct. Plants were sacrificed five days after germination, fixed, embedded in paraffin, and sectioned. Triangles indicate width of meristems. Note that the leaf primordia arising at the flanks of the 35S::LFYmeristem are retarded compared to those on the control meristem. Size bar, 50 ⁇ m.
- the precocious flowers of 35S::LFY tobacco plants are abnormal in organ identity and organ number.
- the floral buds are surrounded by small leaf-like organs, and petals are either absent or sepaloid.
- Stamens and carpels are morphological normal, but their number deviates from wild-type, being in most cases higher. Neither second-order shoots nor flowers develop from the axils of the two true leaves, although adventitious shoots can arise from the hypocotyl.
- pDW151 was introduced into Arabidopsis by vacuum infiltration (Bechtold, et al., C.R. Acad. Sci., 316:1194, 1993). Leaves of adult Arabidopsis thaliana plants of the ecotypes Wassilewskija (Ws-0) and Nossen (No-0) were infiltrated with ASE/pDW151, and seeds were harvested from the infiltrated plants. Seeds were grown on MS medium (Murashige and Skoog, Physiol. Plant 15:473, 1962) supplemented with 50 ⁇ g/ml kanamycin. Transformed plants were identified by their ability to grow on kanamycin containing medium. Using this method, 27 transgenic 35S::LFYArabidopsis plants were isolated, of which 21 exhibited essentially the same dramatic phenotype, which was very similar to that observed in 35S::LFYtobacco plants.
- the transformation experiment utilized a new method that circumvents tissue culture and regeneration of plants from callus, and allows directly for the generation of transgenic seeds (Bechtold, et al., CR. Acad. Sci., 316:1194, 1993).
- leaves of adult plants are vacuum-infiltrated with a suspension of Agrobacterium cells carrying a T-DNA plasmid.
- the Agrobacterium cells grow in planta, where they transfer their T-DNA to host cells, including the precursors of gamete producing cells. Seeds were harvested from the infiltrated plants, and grown on antibiotic containing medium to select for transformants. A small fraction of seeds, between one in several hundred to one in several thousand, were stably transformed with the T-DNA. (A single Arabidopsis plant can produce several thousand seeds.)
- FIG. 4 shows the early flowering phenotype of35S::LFY Arabidopsis plants.
- panel (A) a control plant, transformed with an unrelated construct.
- the rosette leaves (rl) are significantly larger than the cotyledons (cot).
- Panel (B) shows a 35S::LFY transformant (line 151.106).
- the first two rosette leaves (rl) are smaller than the cotyledons. A tiny shoot has formed, with what appear to be two cauline leaves (cl). The floral bud is still unopened. Both plants, which are 17 days old, were selected on kanamycin containing medium for a week, which is likely to have slowed their development somewhat.
- FIG. 5 shows the conversion of all shoots into flowers in 35S::LFY Arabidopsis plants.
- Panel (A) shows a drawing of a mature Arabidopsis plant (Nossen ecotype) of about six weeks of age. Note that indeterminate shoots develop from the axils of all rosette and stem leaves. These shoots bear a few leaves themselves, before they start to produce flowers.
- Panel (B) shows a top view of a wild-type Arabidopsis inflorescence, illustrating the indeterminacy of the shoot meristem. Flowers develop in a phyllotactic spiral, with the youngest flowers being the closest to the center.
- Panels (C)-(E), 35S::LFY plants (generated in the Nossen ecotype), three weeks old.
- Panel (C) Replacement shoots with single flowers (triangles) (line 151.201).
- a cotyledon is indicated (cot).
- Panel (D) Development of a primary terminal flower (1°) on the main shoot, and development of single secondary flower (2°) in the axil of a cauline leaf (cl).
- Single terminal flowers arising from the axils of curled rosette leaves (rl) are indicated by triangles (line 151.209).
- Panel (E) Close-up view of primary and secondary flower shown in (D), at a different angle. Note that the primary terminal flower is abnormal.
- the number of stamens (st) is reduced, and petals and sepals are absent.
- a single first-whorl organ with leaf-, sepal- and carpel-like features is indicated by an asterisk.
- 35S::LFY Arabidopsis plants flower earlier than wild-type plants. There are only two to five rosette leaves, compared to at least eight in wild-type plants, and a stage 12 floral bud can be visible as early as 17 days after germination (FIG. 4). Since it takes two weeks for the development of a stage 12 flower (Smyth, et al., Plant Cell, 2:755, 1990), flowers must initiate within a few days after germination. This is much earlier than in wild type, where the first flowers are initiated only when a plant is about two weeks old. Unlike tobacco, Arabidopsis has an open inflorescence, meaning that the shoot apical meristem remains undifferentiated until the plant dies.
- the 35S::LFY plants not only flower earlier, but their primary axis terminates with a single flower, similar to the tfl mutant phenotype (see FIG. 1).
- ectopic expression of LFY causes transformation of the indeterminate shoot meristem into a determinate floral meristem.
- no normal lateral flowers are formed before the primary terminal flower develops (FIGS. 4B and 5D).
- Additional terminal flowers develop from the axils of leaves in 35S::LFY plants, indicating a transformation of second-order shoot meristems as well (FIGS. 5C and 5D).
- FIG. 5A illustrates the normal architecture of a mature Arabidopsis plant, with indeterminate shoots arising from the axils of all leaves.
- 35S:LFY plants appear to flower faster than any other early flowering mutant that has been described in Arabidopsis, including the embryonicflower (emf) mutant, which appears to skip the rosette phase of vegetative development (Sung, et al., Science, 258:1645-1647, 1992).
- emf mutants Unfortunately, the exact time of flower initiation in emf mutants has not been reported, but the data presented by Sung, et al., supra indicate that flower primordia are not formed before the plant is at least nine days old, making the emf phenotype distinct from the 35S:LFY phenotype. It appears that emf mutants pause after germination, and then proceed directly to the formation of an inflorescence.
- the Dilleniidae are closely related to the Magnoliidae, the most primitive subclass of dicotyledonous plants.
- the Asteridae are the most advanced subclass of dicotyledons (Cronquist, A., An Integrated System of Classification of Flowering Plants, 1981 (New York: Columbia University Press).
- Hybrid aspen was transformed as described previously (Nilsson, O., et al., ibid). Levels of LFY RNA expression were similar to those of 35S::LFY Arabidopsis, as determined by Northern blot analysis. The number of vegetative leaves varied between the different regenerating shoots. Those with a higher number of vegetative leaves formed roots, allowing for transfer to the greenhouse. Individual flowers were removed either from primary transformants that had been transferred to the greenhouse, or from catkins collected in spring 1995 at Carlshem (Umeá, Sweden) from a tree whose age was determined by counting the number of annual rings in a core extracted with an increment borer at 1.5 m above ground level. Flowers were fixed in formaldehyde/acetic acid/ethanol, and destained in ethanol before photography.
- Panel C is a close-up view of solitary male flower that formed in a leaf axil of a seven-month-old 35S::LFY transformant that had been transferred to the greenhouse.
- Panel d shows a close-up view of male flower removed from wild-type catkin shown in panel e.
- Panel e shows a cluster of male catkins of P. tremula, one of the parental species of hybrid aspen, taken from a 15-year-old tree. Red pigment in anthers is apparent. Scale bars: a,b, 5 mm; c, d, 1 mm; e, 20 mm.
- 35S::LFY the transgene is expressed at higher effective levels in than the endogenous gene.
- expression of the endogenous gene in the center of the shoot meristem (which eventually turns into a flower meristem and forms a terminal flower) is relatively low (Kelly, A. J., et al., supra.), and it is conceivable that the other genes, such as AP1, are the primary regulators of flower-meristem-identity in non-transgenic tobacco.
- 35S::LFY was crossed into various mutant backgrounds.
- a 35S::LFY tranformant (line DW151.117, Wassilewskija ecotype) was crossed to ap1-1 (Landberg erecta ecotype)(Irish, F.V. & Hampshire, I.M., Pl Cell 2:741, 1990.).
- Transheterozygote F 1 progeny was either backcrossed to ap1-1 or allowed to self-fertilize.
- the cal genotype of selfed F 2 progeny was determined by polymerase chain reaction (PCR)(Kempin, S.A., et al., Science, 267:552, 1994).
- FIG. 7 shows that 35S::LFY phenotype is partly suppressed by an ap1 mutation.
- Panel a shows five-week-old plants that carry the erecta mutation.
- the 35S::LFY AP1 + plant (left) has no elongated primary shoot.
- a primary shoot is well developed in the 35S::LFY ap1 plant (middle), although the primary shoot still terminates prematurely, and is shorter than that of the non-transgenic ap1 plant (right).
- Panels b-d show a detailed view of 35S::LFY ap1 plants.
- Panel b shows a close-up view of lateral shoot indicated by arrowhead in panel a.
- Panel c shows emerging shoots in the axils of rosette leaves.
- Panel d shows a top view of primary shoot with terminal flower (tf).
- Panels c and d are from a four-week-old plant.
- the ap1 effects are enhanced further by the cal-1 mutation, although there is no qualitative change in the 35S::LFY ap1 phenotype.
- the ap2-1, ap2-2 and ufo-2 mutations caused only additive phenotypes, and did not significantly affect the shoot-to-flower conversion in 35S::LFY plants.
- the apl-1 mutation suppressed the 35S::LFY phenotype to a notable extent, although terminal flowers were still formed (FIG. 7).
- Both the primary and secondary shoots were affected, with the strongest effects being observed in lateral positions (FIG. 7 b,c ).
- the solitary flowers that develop in the axils of rosette leaves of 35S::LFY AP1 + plants become complex shoots with an average of 10 nodes in 35S:LFY apl plants (FIG. 7 c ).
- the present invention shows that constitutive expression of a single flower meristem identity gene, such as LFY or AP1, can induce precocious flower development in plants as diverse as Arabidopsis, an ephermeral weed, and aspen, a perennial tree.
- a single flower meristem identity gene such as LFY or AP1
- LFY or AP1 can induce precocious flower development in plants as diverse as Arabidopsis, an ephermeral weed, and aspen, a perennial tree.
- the results not only contribute to the understanding of flower development and floral induction, they are also likely to be of interest because shorter flowering times lead to shorter generation times, which in turn allows acceleration of breeding programs.
- Modern crop varieties are the result of continued improvement by breeding and two recently developed technologies have made breeding even more important.
- the first technology is molecular mapping, with which genes encoding desirable traits can be rapidly located within the genome.
Abstract
Description
- This application is a continuation of U.S. patent application Ser. No. 09/204,094, which is a continuation of U.S. patent application Ser. No. 08/576,156, now U.S. Pat. No. 5,844,199, which is a continuation-in-part of U.S. patent application Ser. No. 08/360,336, now U.S. Pat. No. 5,637,785, all of which are hereby incorporated by reference in their entirety.
- 1. Field of the Invention
- The invention relates generally to plant genetic engineering, and specifically to novel genetically engineered plants characterized as having a phenotype of early flower meristem development, and methods for producing such plants.
- 2. Description of Related Art
- Most angiosperm species are induced to flower in response to environmental stimuli such as day length and temperature, and internal cues, such as age. Adult organs of flowering plants develop from groups of stem cells called meristems. The identity of a meristem is inferred from structures it produces: vegetative meristems give rise to roots and leaves, inflorescence meristems give rise to flower meristems, and flower meristems give rise to floral organs such as sepals and petals. Not only are meristems capable of generating new meristems of different identity, but their own identity can change during development. For example, a vegetative shoot meristem can be transformed into an inflorescence meristem upon floral induction, and in some species, the inflorescence meristem itself will eventually become a flower meristem. Despite the importance of meristem transitions in plant development, little is known about the underlying mechanisms.
- Following germination, the shoot meristem produces a series of leaf meristems on its flanks. However, once floral induction has occurred, the shoot meristem switches to the production of flower meristems. Flower meristems produce floral organ primordial which develop individually into sepals, petals, stamens or carpels. Thus, flower formation can be thought of as a series of distinct developmental steps, i.e. floral induction, the formation of flower primordia and the production of flower organs. Mutations disrupting each of the steps have been isolated in a variety of species, suggesting that a genetic hierarchy directs the flowering process (see for review, Weigel and Meyerowitz,In Molecular Basis of Morphogenesis (ed. M. Bernfield). 51st Annual Symposium of the Society for Developmental Biology, pp. 93-107, New York, 1993).
- Recently, studies of two distantly related dicotyledons,Arabidopsis thaliana and Antirrhinum majus, led to the identification of three classes of homeotic genes, acting alone or in combination to determine floral organ identity (Bowman, et al., Development, 112:1, 1991; Carpenter and Coen, Genes Devl., 4:1483, 1990; Schwarz-Sommer, et al., Science, 250:931, 1990). Several of these genes are transcription factors whose conserved DNA-binding domain has been designated the MADS box (Schwarz-Sommer, et al., supra).
- Earlier acting genes that control the identity of flower meristems have also been characterized. Flower meristems are derived from inflorescence meristems in both Arabidopsis and Antirrhinum. Two factors that control the development of meristematic cells into flowers are known. In Arabidopsis, the factors are the products of the LEAFY gene (Weigel, et al.,Cell 69:843, 1992) and the APETALA1 gene (Mandel, et al., Nature 360:273, 1992). When either of these genes is inactivated by mutation, structures combining the properties of flowers and inflorescence develop (Weigel, et al., supra; Irish and Sussex, Plant Cell, 2:741, 1990). In Antirrhinum, the homologue of the Arabidopsis LEAFY gene is FLORICAULA (Coen, et al., Cell, 63:1311, 1990) and that of the APETALA1 gene is SQUAMOSA (Huijser, et al., EMBO J, 11:1239, 1992). The latter pair contains MADS box domains.
- LEAFY is expressed very early in floral anlagen and floral primordia, consistent with it having a direct role in establishing floral meristem identity. In the developing floral primordium, LEAFY expression is detected much earlier than expression of the homeotic genes AG and AP3, suggesting that LEAFY plays a role in controlling the expression of floral homeotic genes.
- There is increasing incentive by those working in the field of plant biotechnology to successfully genetically engineer plants, including the major crop varieties. One genetic modification that would be economically desirable would be to accelerate the flowering time of a plant. Induction of flowering is often the limiting factor for growing crop plants. One of the most important factors controlling induction of flowering is day length, which varies seasonally as well as geographically. There is a need to develop a method for controlling and inducing flowering in plants, regardless of the locale or the environmental conditions, thereby allowing production of crops, at any given time. Since most crop products (e.g., seeds, grains, fruits), are derived from flowers, such a method for controlling flowering would be economically invaluable.
- The present invention arose out of the discovery that a genetically modified plant cell could be produced, from which a whole plant can be regenerated which stably incorporates a flower development genetic trait introduced into the plant cell. Specifically, the trait of early flowering can be imparted on a plant by genetic modification according to the method of the invention.
- In a first embodiment, the present invention provides a genetically modified plant comprising at least one heterologous nucleic acid sequence in its genome and characterized as having modulated floral meristem development. Preferably, the plant is genetically modified by introduction of a nucleic acid sequence encoding the LEAFY protein. Alternatively, the plant is genetically modified by transformation with a nucleic acid sequence encoding the LEAFY protein or a nucleic acid sequence encoding the APETALA1 protein, or both. The invention also provides plant cells, plant tissue and seeds derived from the genetically modified plant.
- In a second embodiment, the invention provides a vector(s) for transformation of a plant cell to modulate flower meristem development, wherein said vector(s) comprises a nucleic acid sequence comprising at least one structural gene encoding a protein that modulates flower meristem development, operably associated with a promoter. Preferably, the vector comprises a nucleic acid sequence encoding the LEAFY protein.
- Also provided is a method of producing a genetically modified plant characterized as having modulated flower meristem development. The method comprises contacting a plant cell with a vector(s), comprising a nucleic acid sequence comprising at least one structural gene encoding a protein for modulating flower meristem development, operably associated with a promoter to obtain a transformed plant cell; producing plants from said transformed plant cell; and selecting a plant exhibiting modulated flower meristem development.
- FIG. 1 shows a schematic illustration of pDW139, which is the parental plasmid for construction of 35S::LFY vectors. Open reading frame of LEAFY(LFY) is hatched; 5′ and 3′ untranslated regions are stippled.
- FIG. 2 shows the early flowering phenotype of 35S::LFY tobacco plants. Left, control plant, transformed with an unrelated construct. Middle and right, two independently derived T2 plants carrying a 35S::LFY transgene (lines 146.21, 146.26). Plants are five weeks old.
- FIG. 3 shows precocious enlargement of apical meristem in 35S::LFY tobacco plants. Panel (A), Control, transformed with an unrelated construct. Panel (B), Experimental plant, transformed with a 35S::LFY construct. Size bar, 50 μm.
- FIG. 4 shows the early flowering phenotype of 35S::LFY Arabidopsis plants. Panel (A), Control plant, transformed with an unrelated construct. The rosette leaves (rl) are significantly larger than the cotyledons (cot). Panel (B), 35S::LFY transformant (line 151.106). The first two rosette leaves (rl) are smaller than the cotyledons. A small shoot has formed, with what appear to be two cauline (=stem) leaves (cl).
- FIG. 5 shows the conversion of all shoots into flowers in 35S::LFY Arabidopsis plants. Panel (A), For comparison, a drawing of a mature Arabidopsis plant (Nossen ecotype) of about six weeks of age is shown. Panel (B), Top view of a wild-type Arabidopsis inflorescence, illustrating the indeterminacy of the shoot meristem. Panels (C)-(E) show 35S::LFY plants (generated in the Nossen ecotype), three weeks old. Panel (C), Replacement of shoots with single flowers (triangles) (line 151.201). A cotyledon is indicated (cot). Panel (D), Development of a primary terminal flower (1°) on the main shoot, and development of single secondary flower (2°) in the axil of a cauline leaf (cl). Single terminal flowers arising from the axils of curled rosette leaves (rl) are indicated by triangles (line 151.209). Panel (E), Close-up view of primary and secondary flower shown in (D), at a different angle. The gynoecium (g), comprising the carpels, appears largely normal. The number of stamens (st) is reduced, and petals and sepals are absent. A single first-whorl organ with leaf-, sepal- and carpel-like features is indicated by an asterisk.
- FIG. 6 shows constitutive expression of Arabidopsis LFY converts aspen shoots into flowers. Panels a and b show five-month-old shoots of hybrid aspen (Populous tremula x tremuloides) grown in tissue culture. Panel a shows a 35S::LFY transformant. Solitary, lateral flowers in the axils of leaves (lf) and an abnormal terminal flower (tf) are indicated. Panel b shows a non-transgenic control. Arrowheads indicate axils of leaves, from which lateral vegetative shoots will emerge, normally in the following year. Note that aspen plants regenerated from tissue culture show the same juvenile phenotype during the first growing cycle as plants grown from seed (Nilsson, O., Thesis, Swedish Univ. Agricul. Sciences, 1995) Panel C is a close-up view of solitary male flower that formed in a leaf axil of a seven-month-old 35S::LFY transformant that had been transferred to the greenhouse. Panel d shows a close-up view of male flower removed from wild-type catkin shown in panel e. Note bract (b) subtending wild-type flower. Panel e shows a cluster of male catkins of P. tremula, one of the parental species of hybrid aspen, taken from a 15-year-old tree. Red pigment in anthers is apparent. Scale bars: a, b, 5 mm; c, d, 1 mm; e, 20 mm.
- FIG. 7 shows 35S::LFY phenotype is partly suppressed by an ap1 mutation. Panel a shows five-week-old plants that carry the erecta mutation. The 35S::LFY AP1+plant (left) has no elongated primary shoot. A primary shoot is well developed in the 35S::LFY ap1 plant (middle), although the primary shoot still terminates prematurely, and is shorter than that of the non-transgenic ap1 plant (right). Panels b-d show a detailed view of 35S::LFY ap1 plants. Panel b shows a close-up view of lateral shoot indicated by arrowhead in panel a. Panel c shows emerging shoots in the axils of rosette leaves. Panel d shows a top view of primary shoot with terminal flower (tf). Panels c and d are from a four-week-old plant. The ap1 effects are enhanced further by the cal-1 mutation, although there is no qualitative change in the 35S::LFY ap1 phenotype.
- The present invention provides a genetically modified plant which is characterized as having the phenotypic trait of early flower development, or early flowering. The plant is genetically modified by at least one structural gene that encodes a protein, such as LEAFY, which is sufficient to induce flowering in the plant.
- In a first embodiment, the invention provides a genetically modified plant comprising at least one heterologous nucleic acid sequence in its genome and characterized as having modulated flower meristem development. Also included herein are plant cells and plant tissue, all derived from the genetically modified plant of the invention. In addition, seeds which can germinate into a genetically modified plant as described herein are also provided.
- The term “genetic modification” as used herein refers to the introduction of one or more heterologous nucleic acid sequences into one or more plant cells, which can generate whole, sexually competent, viable plants. The term “genetically modified” as used herein refers to a plant which has been generated through the aforementioned process. Genetically modified plants of the invention are capable of self-pollinating or cross-pollinating with other plants of the same species so that the foreign gene, carried in the germ line, can be inserted into or bred into agriculturally useful plant varieties. The term “plant cell” as used herein refers to protoplasts, gamete producing cells, and cells which regenerate into whole plants. Accordingly, a seed comprising multiple plant cells capable of regenerating into a whole plant, is included in the definition of “plant cell”.
- As used herein, the term “plant” refers to either a whole plant, a plant part, a plant cell, or a group of plant cells, such as plant tissue, for example. Plantlets are also included within the meaning of “plant”. Plants included in the invention are any flowering plants amenable to transformation techniques, including both monocotyledonous and dicotyledonous plants.
- Examples of monocotyledonous plants include, but are not limited to, asparagus, field and sweet corn, barley, wheat, rice, sorghum, onion, pearl millet, rye and oats. Examples of dicotyledonous plants include, but are not limited to tomato, tobacco, cotton, rapeseed, field beans, soybeans, peppers, lettuce, peas, alfalfa, clover, cole crops orBrassica oleracea (e.g., cabbage, broccoli, cauliflower, brussel sprouts), radish, carrot, beets, eggplant, spinach, cucumber, squash, melons, cantaloupe, sunflowers and various ornamentals. Exemplary models described herein include the tobacco plant and the perennial tree, aspen.
- The term “heterologous nucleic acid sequence” as used herein refers to at least one structural gene operably associated with a regulatory sequence such as a promoter. The nucleic acid sequence originates in a foreign species, or, in the same species if substantially modified from its original form. For example, the term “heterologous nucleic acid sequence” includes a nucleic acid originating in the same species, where such sequence is operably linked to a promoter that differs from the natural or wild-type promoter.
- As used herein, the term “nucleic acid sequence” refers to a polymer of deoxyribonucleotides or ribonucleotides, in the form of a separate fragment or as a component of a larger construct. DNA encoding the proteins utilized in the method of the invention can be assembled from cDNA fragments or from oligonucleotides which provide a synthetic gene which is capable of being expressed in a recombinant transcriptional unit. Polynucleotide or nucleic acid sequences of the invention include DNA, RNA and cDNA sequences.
- Examples of structural genes that may be employed in the present invention include the LEAFY gene and the APETALA1 gene which control flowering. Also included in the present invention are structural and functional homologues of the LEAFY and APETALA1 genes. For example, inAntirrhinum majus, the snapdragon, the homologue of the LEAFY gene is the FLORICAULA gene and the homologue of the APETALA1 gene is the SQUAMOSA gene. Other genes which control flowering will be known to those of skill in the art or can be readily ascertained.
- Nucleic acid sequences utilized in the invention can be obtained by several methods. For example, the DNA can be isolated using hybridization procedures which are well known in the art. These include, but are not limited to: 1) hybridization of probes to genomic or cDNA libraries to detect shared nucleotide sequences; 2) antibody screening of expression libraries to detect shared structural features and 3) synthesis by the polymerase chain reaction (PCR). Sequences for specific genes can also be found in GenBank, National Institutes of Health computer database.
- Hybridization procedures useful for screening for desired nucleic acid sequences utilized herein employ labeled mixed synthetic oligonucleotide probes where each probe is potentially the complete complement of a specific DNA sequence in the hybridization sample which includes a heterogeneous mixture of denatured double-stranded DNA. For such screening, hybridization is preferably performed on either single-stranded DNA or denatured double-stranded DNA. Hybridization is particularly useful in the detection of cDNA clones derived from sources where an extremely low amount of mRNA sequences relating to the polypeptide of interest are present. In other words, by using stringent hybridization conditions directed to avoid non-specific binding, it is possible, for example, to allow the autoradiographic visualization of a specific cDNA clone by the hybridization of the target DNA to that single probe in the mixture which is its complete complement (Wallace, et al.,Nucleic Acid Research, 9:879, 1981).
- Specific DNA sequences encoding a heterologous protein of interest, such as LEAFY protein, can also be obtained by: 1) isolation of double-stranded DNA sequences from the genomic DNA; 2) chemical synthesis of a DNA sequence to provide the necessary codons for the polypeptide of interest; and 3) in vitro synthesis of a double-stranded DNA sequence by reverse transcription of mRNA isolated from a eukaryotic donor cell. In the latter case, a double-stranded DNA complement of mRNA is eventually formed which is generally referred to as cDNA.
- A cDNA expression library, such as lambda gtl 11, can be screened indirectly for a heterologous polypeptide having at least one epitope, using antibodies specific for the heterologous protein. Such antibodies can be either polyclonally or monoclonally derived and used to detect expression product indicative of the presence of heterologous protein cDNA.
- A polypeptide sequence can be deduced from the genetic code, however, the degeneracy of the code must be taken into account. Nucleic acid sequences utilized in the invention include sequences which are degenerate as a result of the genetic code. There are 20 natural amino acids, most of which are specified by more than one codon. Therefore, as long as the amino acid sequence of heterologous polypeptide results in a functional polypeptide (at least, in the case of the sense polynucleotide strand), all degenerate nucleotide sequences are included in the invention.
- “Modulated” flower meristem development as used herein, refers to flower development in the plant which may be either accelerated or inhibited/delayed as compared to the naturally occurring, unmanipulated plant. Therefore, the term “modulate” envisions the acceleration or augmentation of flower development when development is desirable or suppression or inhibition of flower development when development is not desirable.
- The heterologous nucleic acid sequences utilized herein are structural genes for flower meristem development. Preferably, such genes encode a protein that is sufficient for the initiation of flowering, and most preferably, the nucleic acid sequence encodes the LEAFY protein. The LEAFY gene or other flower meristem development gene may be utilized alone or in combination with another structural gene, such as another gene which encodes a protein important in the development of flowering. An example of such a gene is the APETALA1 gene.
- Genetically modified plants of the present are produced by contacting a plant cell with a vector comprising a heterologous nucleic acid sequence comprising at least one structural gene encoding a protein that modulates flower meristem development. To be effective once introduced into plant cells, the structural gene of interest must be operably associated with a promoter which is effective in the plant cells to cause transcription of the gene of interest. Additionally, a polyadenylation sequence or transcription control sequence, also recognized in plant cells may also be employed. It is preferred that the vector harboring the heterologous nucleic acid sequence also contain one or more selectable marker genes so that the transformed cells can be selected from non-transformed cells in culture, as described herein.
- The term “operably associated” refers to functional linkage between a promoter sequence and the structural gene regulated by the promoter nucleic acid sequence. The operably linked promoter controls the expression of the polypeptide encoded by the structural gene.
- The expression of structural genes employed in the present invention may be driven by a number of promoters. Although the endogenous promoter of a structural gene of interest may be utilized for transcriptional regulation of the gene, preferably, the promoter is a foreign regulatory sequence. For plant expression vectors, suitable viral promoters include the 35S RNA and 19S RNA promoters of CaMV (Brisson, et al.,Nature, 310:511, 1984; Odell, et al., Nature, 313:810, 1985); the full-length transcript promoter from Figwort Mosaic Virs (FMV) (Gowda, et al., J Cell Biochem., 13D: 301, 1989) and the coat protein promoter to TMV (Takamatsu, et al., EMBO J. 6:307, 1987). Alternatively, plant promoters such as the light-inducible promoter from the small subunit of ribulose bis-phosphate carboxylase (ssRUBISCO) (Coruzzi, et al., EMBO J., 3:1671, 1984; Broglie, et al., Science, 224:838, 1984); mannopine synthase promoter (Velten, et al., EMBO J., 3:2723, 1984) nopaline synthase (NOS) and octopine synthase (OCS) promoters (carried on tumor-inducing plasmids ofAgrobacterium tumefaciens) or heat shock promoters, e.g., soybean hsp 17.5-E or hsp 17.3-B (Gurley, et al., Mol. Cell. Biol., 6:559, 1986; Severin, et al., Plant Mol. Biol., 15:827, 1990) may be used.
- Promoters useful in the invention include both constitutive and inducible natural promoters as well as engineered promoters. The CaMV promoters are examples of constitutive promoters. To be most useful, an inducible promoter should 1) provide low expression in the absence of the inducer; 2) provide high expression in the presence of the inducer; 3) use an induction scheme that does not interfere with the normal physiology of the plant; and 4) have no effect on the expression of other genes. Examples of inducible promoters useful in plants include those induced by chemical means, such as the yeast metallothionein promoter which is activated by copper ions (Mett, et al.,Proc. Nat. Acad. Sci., U.S.A., 90:4567, 1993); In2-1 and In2-2 regulator sequences which are activated by substituted benzenesulfonamides, e.g., herbicide safeners (Hershey, et al., Plant Mol. Biol., 17:679, 1991); and the GRE regulatory sequences which are induced by glucocorticoids (Schena, et al., Proc. Natl. Acad. Sci., USA., 88:10421, 1991). Other promoters, both constitutive and inducible and enhancers will be known to those of skill in the art.
- The particular promoter selected should be capable of causing sufficient expression to result in the production of an effective amount of the structural gene product, e.g., LEAFY, to cause early floral meristem development: The promoters used in the vector constructs of the present invention may be modified, if desired, to affect their control characteristics.
- Tissue specific promoters may also be utilized in the present invention. An example of a tissue specific promoter is the promoter expressed in shoot meristems (Atanassova, et al.,Plant J., 2:291, 1992). Other tissue specific promoters useful in transgenic plants, including the cdc2a promoter and cyc07 promoter, will be known to those of skill in the art. (See for example, Ito, et al., Plant Mol. Biol., 24:863, 1994; Martinez, et al., Proc. Natl. Acad. Sci. USA, 89:7360, 1992; Medford, et al., Plant Cell, 3:359, 1991; Terada, et al., Plant Journal, 3:241, 1993; Wissenbach, et al., Plant Journal, 4:411, 1993).
- Optionally, a selectable marker may be associated with the heterologous nucleic acid sequence, i.e., the structural gene operably linked to a promoter. As used herein, the term “marker” refers to a gene encoding a trait or a phenotype which permits the selection of, or the screening for, a plant or plant cell containing the marker. Preferably, the marker gene is an antibiotic resistance gene whereby the appropriate antibiotic can be used to select for transformed cells from among cells that are not transformed. Examples of suitable selectable markers include adenosine deaminase, dihydrofolate reductase, hygromycin-B-phosphotransferase, thymidine kinase, xanthine-guanine phospho-ribosyltransferase and amino-
glycoside 3′-O-phosphotransferase II (kanamycin, neomycin and G418 resistance). Other suitable markers will be known to those of skill in the art. - Vector(s) employed in the present invention for transformation of a plant cell to modulate flower meristem development comprise a nucleic acid sequence comprising at least one structural gene encoding a protein that modulates flower meristem development, operably associated with a promoter. To commence a transformation process in accordance with the present invention, it is first necessary to construct a suitable vector and properly introduce it into the plant cell. The details of the construction of the vectors then utilized herein are known to those skilled in the art of plant genetic engineering.
- For example, the heterologous nucleic acid sequences utilized in the present invention can be introduced into plant cells using Ti plasmids, root-inducing (Ri) plasmids, and plant virus vectors. (For reviews of such techniques see, for example, Weissbach & Weissbach, 1988, Methods for Plant Molecular Biology, Academic Press, NY, Section VIII, pp. 421-463; and Grierson & Corey, 1988, Plant Molecular Biology, 2d Ed., Blackie, London, Ch. 7-9, and Horsch, et al.,Science, 227:1229, 1985, both incorporated herein by reference).
- One of skill in the art will be able to select an appropriate vector for introducing the heterologous nucleic acid sequence in a relatively intact state. Thus, any vector which will produce a plant carrying the introduced DNA sequence should be sufficient. Even a naked piece of DNA would be expected to be able to confer the properties of this invention, though at low efficiency. The selection of the vector, or whether to use a vector, is typically guided by the method of transformation selected.
- The transformation of plants in accordance with the invention may be carried out in essentially any of the various ways known to those skilled in the art of plant molecular biology. (See, for example, Methods of Enzymology, Vol. 153, 1987, Wu and Grossman, Eds., Academic Press, incorporated herein by reference). As used herein, the term “transformation” means alteration of the genotype of a host plant by the introduction of a heterologous nucleic acid sequence.
- For example, a heterologous nucleic acid sequence can be introduced into a plant cell utilizingAgrobacterium tumefaciens containing the Ti plasmid. In using an A. tumefaciens culture as a transformation vehicle, it is most advantageous to use a non-oncogenic strain of the Agrobacterium as the vector carrier so that normal non-oncogenic differentiation of the transformed tissues is possible. It is also preferred that the Agrobacterium harbor a binary Ti plasmid system. Such a binary system comprises 1) a first Ti plasmid having a virulence region essential for the introduction of transfer DNA (T-DNA) into plants, and 2) a chimeric plasmid. The latter contains at least one border region of the T-DNA region of a wild-type Ti plasmid flanking the nucleic acid to be transferred. Binary Ti plasmid systems have been shown effective to transform plant cells (De Framond, Biotechnology, 1:262, 1983; Hoekema, et al., Nature, 303:179, 1983). Such a binary system is preferred because it does not require integration into Ti plasmid in Agrobacterium.
- Methods involving the use of Agrobacterium include, but are not limited to: 1) co-cultivation of Agrobacterium with cultured isolated protoplasts; 2) transformation of plant cells or tissues with Agrobacterium; or 3) transformation of seeds, apices or meristems with Agrobacterium.
- In addition, gene transfer can be accomplished by in situ transformation by Agrobacterium, as described by Bechtold, et al., (C.R. Acad. Sci. Paris, 316:1194, 1993) and exemplified in the Examples herein. This approach is based on the vacuum infiltration of a suspension of Agrobacterium cells.
- The preferred method of introducing heterologous nucleic acid into plant cells is to infect such plant cells, an explant, a meristem or a seed, with transformedAgrobacterium tumefaciens as described above. Under appropriate conditions known in the art, the transformed plant cells are grown to form shoots, roots, and develop further into plants. A preferred vector(s) of the invention comprises a Ti plasmid binary system wherein the heterologous nucleic acid sequence encodes the LEAFY protein. Such a vector may optionally contain a nucleic acid sequence which encodes a second flower development factor, such as APETALA1. Alternatively, two vectors can be utilized wherein each vector contains a heterologous nucleic acid sequence. Other flower development genes can be utilized for construction of one or more vectors, in a similar manner.
- Alternatively, heterologous nucleic acid can be introduced into a plant cell by contacting the plant cell using mechanical or chemical means. For example, the nucleic acid can be mechanically transferred by microinjection directly into plant cells by use of micropipettes. Alternatively, the nucleic acid may be transferred into the plant cell by using polyethylene glycol which forms a precipitation complex with genetic material that is taken up by the cell.
- Heterologous nucleic acid can also be introduced into plant cells by electroporation (Fromm, et al,Proc. Natl. Acad. Sci., U.S.A., 82:5824, 1985, which is incorporated herein by reference). In this technique, plant protoplasts are electroporated in the presence of vectors or nucleic acids containing the relevant nucleic acid sequences. Electrical impulses of high field strength reversibly permeabilize membranes allowing the introduction of nucleic acids. Electroporated plant protoplasts reform the cell wall, divide and form a plant callus. Selection of the transformed plant cells with the transformed gene can be accomplished using phenotypic markers as described herein.
- Another method for introducing nucleic acid into a plant cell is high velocity ballistic penetration by small particles with the nucleic acid to be introduced contained either within the matrix of small beads or particles, or on the surface thereof (Klein, et al.,Nature 327:70, 1987). Although, typically only a single introduction of a new nucleic acid sequence is required, this method particularly provides for multiple introductions.
- Cauliflower mosaic virus (CaMV) may also be used as a vector for introducing heterologous nucleic acid into plant cells (U.S. Pat. No. 4,407,956). CaMV viral DNA genome is inserted into a parent bacterial plasmid creating a recombinant DNA molecule which can be propagated in bacteria. After cloning, the recombinant plasmid again may be cloned and further modified by introduction of the desired nucleic acid sequence. The modified viral portion of the recombinant plasmid is then excised from the parent bacterial plasmid, and used to inoculate the plant cells or plants.
- In another embodiment, the invention includes a method of producing a genetically modified plant characterized as having modulated flower meristem development, said method comprising contacting a plant cell with a vector, comprising a heterologous nucleic acid sequence comprising at least one structural gene encoding a protein for modulating flower meristem development, operably associated with a promoter to obtain a transformed plant cell; growing a plant from said transformed plant cell; and selecting a plant exhibiting modulated flower meristem development.
- As used herein, the term “contacting” refers to any means of introducing the vector(s) into the plant cell, including chemical and physical means as described above. Preferably, contacting refers to introducing the nucleic acid or vector into plant cells (including an explant, a meristem or a seed), viaAgrobacterium tumefaciens transformed with the heterologous nucleic acid as described above.
- Normally, a plant cell is regenerated to obtain a whole plant from the transformation process. The immediate product of the transformation is referred to as a “transgenote”. The term “growing” or “regeneration” as used herein means growing a whole plant from a plant cell, a group of plant cells, a plant part (including seeds), or a plant piece (e.g., from a protoplast, callus, or tissue part).
- Regeneration from protoplasts varies from species to species of plants, but generally a suspension of protoplasts is first made. In certain species, embryo formation can then be induced from the protoplast suspension, to the stage of ripening and germination as natural embryos. The culture media will generally contain various amino acids and hormones, necessary for growth and regeneration. Examples of hormones utilized include auxin and cytokinins. It is sometimes advantageous to add glutamic acid and proline to the medium, especially for such species as corn and alfalfa. Efficient regeneration will depend on the medium, on the genotype, and on the history of the culture. If these variables are controlled, regeneration is reproducible.
- Regeneration also occurs from plant callus, explants, organs or parts. Transformation can be performed in the context of organ or plant part regeneration. (seeMethods in Enzymology, Vol. 118 and Klee, et al., Annual Review of Plant Physiology, 38:467, 1987). Utilizing the leaf disk-transformation-regeneration method of Horsch, et al., Science, 227:1229, 1985, disks are cultured on selective media, followed by shoot formation in about 2-4 weeks. Shoots that develop are excised from calli and transplanted to appropriate root-inducing selective medium. Rooted plantlets are transplanted to soil as soon as possible after roots appear. The plantlets can be repotted as required, until reaching maturity.
- In vegetatively propagated crops, the mature transgenic plants are propagated by the taking of cuttings or by tissue culture techniques to produce multiple identical plants. Selection of desirable transgenotes is made and new varieties are obtained and propagated vegetatively for commercial use. In seed propagated crops, the mature transgenic plants can be self crossed to produce a homozygous inbred plant. The inbred plant produces seed containing the newly introduced foreign gene(s). These seeds can be grown to produce plants that would produce the selected phenotype, e.g. early flowering.
- Parts obtained from the regenerated plant, such as flowers, seeds, leaves, branches, fruit, and the like are included in the invention, provided that these parts comprise cells that have been transformed as described. Progeny and variants, and mutants of the regenerated plants are also included within the scope of the invention, provided that these parts comprise the introduced nucleic acid sequences.
- Plants exhibiting modulated flower meristem development can be selected by visual observation. The invention includes a plant produced by the method of the invention, including plant tissue, seeds, and other plant cells derived from the genetically modified plant.
- In yet another embodiment, the invention provides a method for modulating flower meristem development in a plant cell, said method comprising contacting said plant cell with a vector as described above to obtain a transformed plant cell, growing the transformed plant cell under plant forming conditions, and modulating flower meristem development in the plant. The method of the invention requires that the promoter sequence operably linked with the structural gene. The promoter is an inducible promoter when induction of flower development is desired. For example, a plant cell and plant is produced as described above and modulated flower meristem development is induced by contacting the promoter, linked with a nucleic acid sequence encoding LEAFY, with an appropriate inducer. Such inducible promoters are described above, and include those promoters preferably inducible by chemical means.
- While the present examples demonstrate that constitutive expression of a floral regulatory gene (LEAFY) causes accelerated flowering, this system could be modified such that flowering would be inhibited. For example, dominant-negative versions of floral regulatory genes could be expressed constitutively. Dominant-negative mutants are proteins that actively interfere with the function of a normal, endogenous protein. Thus, the action of a gene can be blocked without inactivating the structural gene itself or its RNA. This strategy has been successful for transcription factors (e.g., Attardi, et al.,Proc. Natl. Acad Sci. USA, 90:10563, 1993; Lloyd, et al., Nature, 352:635, 1991; Logeat, et al., EMBO J., 10:1827, 1991: Mantovani, et al., J. Biol. Chem., 269:20340, 1994; Ransone, et al., Proc. Natl. Acad. Sci. USA, 87:3806, 1990; Richardson, et al., Mech. Dev., 45:173, 1994; Tsai, et al., Genes Dev., 6:2258, 1992.) The LEAFY protein is likely to be a transcription factor, as it localizes to the nucleus and can bind to DNA in vitro. Likewise, most other floral regulatory genes, including APETALA1, encode known transcription factors with a MADS DNA-binding domain (e.g., Mandel, et al., Nature, 360:273, 1992).
- The above disclosure generally describes the present invention. A more complete understanding can be obtained by reference to the following specific examples which are provided herein for purposes of illustration only and are not intended to limit the scope of the invention.
- To study the effects of ectopic LEAFY (LFY) expression, a chimeric gene in which the LFY coding region is under the control of the constitutive 35S promoter from cauliflower mosaic virus (CaMV) was constructed (Odell, J.T., et al.Nature, 313:810, 1985.) By way of illustration, the chimeric 35S::LFY gene was introduced into Arabidopsis and the distantly related tobacco plants by T-DNA mediated transformation. Other examples show aspen trees transformed with LFY. The phenotypic effects observed in transgenic plants show that LFY is not only necessary, but also sufficient for the initiation of flower development.
- For transformation of tobacco, the pDW146 vector was used. For transformation of Arabidopsis, the pDW151 vector was used. Both vectors are derived from plasmid pDW139, which contains the entire open reading frame of the LEAFY (LFY) gene fromArabidopsis thaliana (Weigel, D., et al., Cell, 69:843, 1992), plus 21 bp upstream of the initiation codon and 195 bp downstream of the stop codon (for cDNA sequence, see Weigel et al., supra; genomic sequence deposited in GenBank under accession number M91208). At the 5′ end, a Bg12 site was added by polymerase chain reaction (Saiki, et al., Science, 239:487, 1988). At the 3′ end, a genomic ScaI site was eliminated in the cloning process, and it is followed immediately by an Asp718 site derived from the pBluescript KS+cloning vector (FIG. 1).
- FIG. 1 shows a schematic illustration of pDW139 parental plasmid for construction of 35S::LFY vectors. The open reading frame of LEAFY (LFY) is hatched; 5′ and 3′ untranslated regions are stippled.
- To construct pDW146, the 1.5 kb Bg12/Asp718 fragment carrying the LFY sequences was inserted into the binary T-DNA transformation vector pMON530 (Rogers, et al.,Meth. Enzymol, 153:253, 1987), using the same sites in the vector. This vector contains an expression cassette comprising a 0.3 kb fragment of the cauliflower mosaic virus 35S promoter, including the transcription initiation site (Guilley, et al., Cell, 30:763, 1982); Odell, et al., supra.); a multilinker containing several unique restriction sites; and a functional polyadenylation signal from the Ti plasmid T-DNA nopaline synthase gene (“3′ nos”; [Bevan, et al., Plant Cell, 1:141, 1983]).
- To construct pDW151, the Asp718 site of pDW139 was filled in with Klenow enzyme (Sambrook, et al.,Molecular Cloning 2nd ed. (Cold Spring Harbor: Cold Spring Harbor Laboratory, 1989) and a Bg12 linker was added. The resulting Bg12 fragment was inserted into the BamH1 site of pCGN18, a transformation vector containing a
CaMV 35S promoter 3′ nos expression cassette (Jack, et al, Cell, 76:703, 1994). - pDW146 and pDW151 plasmid DNAs isolated fromE. coli were transformed into Agrobacterium tumefaciens strain LBA4404 (Ooms, et al, Plasmid, 7:15, 1982) or ASE (Fraley, et al, Biotechnology 3:629, 1985), respectively, using the freeze-thaw method as described (Hofgen and Willmitzer, Nucl. Acids Res, 16:9877, 1988), except that LB medium (Sambrook, et al., supra) was used instead of YEB.
- For generation of transgenic tobacco plants, leaf pieces of sterily grown tobacco strainNicotiana tabacum var. Xanthi were infected with LBA4404/pDW146, and plants were regenerated as described (Horsch, et al., Science, 227:1229, 1985). Selection for transformed plants was with 200 μg/ml kanamycin. Kanamycin resistant regenerated plants were transferred to soil, and seeds were harvested from the primary transformants.
- FIG. 2 shows the early flowering phenotype of 35S::LFY tobacco plants. The left panel shows a control plant, transformed with an unrelated construct (a LFY promoter fused to a GUS reporter gene). The middle and right panels show two independently derived T2 plants carrying a 35S::LFY transgene (lines 146.21, 146.26). The plants shown are five weeks old. Note abundant proliferation of leaves in the control, while the experimental plants have produced only two true leaves before initiating a terminal flower. The insert shows a top view of floral bud of plant shown at the right. The bud is still unopened.
- Of 32 transgenic tobacco lines analyzed in detail, 27 exhibit the same dramatic phenotype in the progeny of the primary transformants (T2 generation). Transgenic plants develop only one pair of true leaves, in addition to the embryonic leaves (cotyledons), before they produce a terminal flower (FIG. 2). Although wild-type tobacco plants also produce a terminal flower, they generate twenty to twenty-five pairs of leaves before flowering. Thus, constitutive LFY expression causes precocious conversion of the shoot meristem into a floral meristem. Histological sections of transgenic plants reveal that the apical meristem is morphologically different from that of untransformed plants at least as early as five days after germination (FIG. 3). The result of these changes is that transformed plants produce visible floral buds after two weeks, while normal tobacco plants flower only after about three to five months (the exact time depends on environmental conditions, such as light intensity, fertilizer, size of pots in which plants are grown, etc.).
- FIG. 3 shows precocious enlargement of apical meristem in 35S::LFY tobacco plants. Panel (A) is a control, transformed with the unrelated construct described in FIG. 2. Panel (B) shows an experimental plant, transformed with a 35S::LFY construct. Plants were sacrificed five days after germination, fixed, embedded in paraffin, and sectioned. Triangles indicate width of meristems. Note that the leaf primordia arising at the flanks of the 35S::LFYmeristem are retarded compared to those on the control meristem. Size bar, 50 μm.
- The precocious flowers of 35S::LFY tobacco plants are abnormal in organ identity and organ number. The floral buds are surrounded by small leaf-like organs, and petals are either absent or sepaloid. Stamens and carpels are morphological normal, but their number deviates from wild-type, being in most cases higher. Neither second-order shoots nor flowers develop from the axils of the two true leaves, although adventitious shoots can arise from the hypocotyl.
- pDW151 was introduced into Arabidopsis by vacuum infiltration (Bechtold, et al.,C.R. Acad. Sci., 316:1194, 1993). Leaves of adult Arabidopsis thaliana plants of the ecotypes Wassilewskija (Ws-0) and Nossen (No-0) were infiltrated with ASE/pDW151, and seeds were harvested from the infiltrated plants. Seeds were grown on MS medium (Murashige and Skoog, Physiol. Plant 15:473, 1962) supplemented with 50 μg/ml kanamycin. Transformed plants were identified by their ability to grow on kanamycin containing medium. Using this method, 27 transgenic 35S::LFYArabidopsis plants were isolated, of which 21 exhibited essentially the same dramatic phenotype, which was very similar to that observed in 35S::LFYtobacco plants.
- The transformation experiment utilized a new method that circumvents tissue culture and regeneration of plants from callus, and allows directly for the generation of transgenic seeds (Bechtold, et al.,CR. Acad. Sci., 316:1194, 1993). In this method, leaves of adult plants are vacuum-infiltrated with a suspension of Agrobacterium cells carrying a T-DNA plasmid. The Agrobacterium cells grow in planta, where they transfer their T-DNA to host cells, including the precursors of gamete producing cells. Seeds were harvested from the infiltrated plants, and grown on antibiotic containing medium to select for transformants. A small fraction of seeds, between one in several hundred to one in several thousand, were stably transformed with the T-DNA. (A single Arabidopsis plant can produce several thousand seeds.)
- The following description of the 35S::LFY phenotype in Arabidopsis is based on the analysis of first generation transformants. The phenotype should not change significantly in subsequent generations, because these transformants have been grown from seeds, as opposed to having been regenerated from tissue culture. The same method has been used to generate transformants with four other constructs, none of which cause the phenotype observed with the 35 S::LFY construct. FIG. 4 shows the early flowering phenotype of35S::LFY Arabidopsis plants. In panel (A), a control plant, transformed with an unrelated construct. The rosette leaves (rl) are significantly larger than the cotyledons (cot). Panel (B) shows a 35S::LFY transformant (line 151.106). The first two rosette leaves (rl) are smaller than the cotyledons. A tiny shoot has formed, with what appear to be two cauline leaves (cl). The floral bud is still unopened. Both plants, which are 17 days old, were selected on kanamycin containing medium for a week, which is likely to have slowed their development somewhat.
- FIG. 5 shows the conversion of all shoots into flowers in 35S::LFY Arabidopsis plants. Panel (A) shows a drawing of a mature Arabidopsis plant (Nossen ecotype) of about six weeks of age. Note that indeterminate shoots develop from the axils of all rosette and stem leaves. These shoots bear a few leaves themselves, before they start to produce flowers. Panel (B) shows a top view of a wild-type Arabidopsis inflorescence, illustrating the indeterminacy of the shoot meristem. Flowers develop in a phyllotactic spiral, with the youngest flowers being the closest to the center. Panels (C)-(E), 35S::LFY plants (generated in the Nossen ecotype), three weeks old. Panel (C), Replacement shoots with single flowers (triangles) (line 151.201). A cotyledon is indicated (cot). Panel (D), Development of a primary terminal flower (1°) on the main shoot, and development of single secondary flower (2°) in the axil of a cauline leaf (cl). Single terminal flowers arising from the axils of curled rosette leaves (rl) are indicated by triangles (line 151.209). Panel (E), Close-up view of primary and secondary flower shown in (D), at a different angle. Note that the primary terminal flower is abnormal. The gynoecium (g), comprising the carpels, appears largely normal. The number of stamens (st) is reduced, and petals and sepals are absent. A single first-whorl organ with leaf-, sepal- and carpel-like features is indicated by an asterisk.
- 35S::LFY Arabidopsis plants flower earlier than wild-type plants. There are only two to five rosette leaves, compared to at least eight in wild-type plants, and a stage 12 floral bud can be visible as early as 17 days after germination (FIG. 4). Since it takes two weeks for the development of a stage 12 flower (Smyth, et al.,Plant Cell, 2:755, 1990), flowers must initiate within a few days after germination. This is much earlier than in wild type, where the first flowers are initiated only when a plant is about two weeks old. Unlike tobacco, Arabidopsis has an open inflorescence, meaning that the shoot apical meristem remains undifferentiated until the plant dies. The 35S::LFY plants not only flower earlier, but their primary axis terminates with a single flower, similar to the tfl mutant phenotype (see FIG. 1). Thus, ectopic expression of LFY causes transformation of the indeterminate shoot meristem into a determinate floral meristem. In contrast with the tfl mutant phenotype, no normal lateral flowers are formed before the primary terminal flower develops (FIGS. 4B and 5D). Additional terminal flowers develop from the axils of leaves in 35S::LFY plants, indicating a transformation of second-order shoot meristems as well (FIGS. 5C and 5D). Surprisingly, most 35S::LFY plants develop a tiny shoot, with one or two leaves that resemble cauline (stem) leaves of wild type (FIGS. 4B and 5D). For comparison, FIG. 5A illustrates the normal architecture of a mature Arabidopsis plant, with indeterminate shoots arising from the axils of all leaves.
- The early flowering phenotype, and the transformation of a shoot into a floral meristem, show that LFY activity is sufficient to determine the identity of a meristem. However, since the shoot meristem produces leaves before it is converted into a floral meristem, there appear to be additional factors that prevent the shoot meristem from responding to LFY activity immediately after germination.
- 35S:LFY plants appear to flower faster than any other early flowering mutant that has been described in Arabidopsis, including the embryonicflower (emf) mutant, which appears to skip the rosette phase of vegetative development (Sung, et al.,Science, 258:1645-1647, 1992). Unfortunately, the exact time of flower initiation in emf mutants has not been reported, but the data presented by Sung, et al., supra indicate that flower primordia are not formed before the plant is at least nine days old, making the emf phenotype distinct from the 35S:LFY phenotype. It appears that emf mutants pause after germination, and then proceed directly to the formation of an inflorescence.
- The exact phenotype of individual 35S::LFY Arabidopsis plants varies. Most flowers observed are virtually identical to wild-type flowers (FIGS. 5D and E). Very importantly, for further analysis, stamens and carpels are fertile. The primary terminal flowers are often abnormal, in that the outer organs are leaf-like or absent, and the numbers of petals is reduced, similar to the effect seen in the terminal flowers of 35S::LFY tobacco plants (FIG. 5E). In addition, carpels can be unfused, and the number of stamens can be lower than the wild-type number of six.
- The finding that the LEAFY gene from Arabidopsis can modify flowering in tobacco implies that the mode of LEAFY function is well conserved among flowering plants, that the Arabidopsis gene is likely to function in a wide variety of flowering plants. Arabidopsis and tobacco belong to two very divergent subclasses among the class of dicotyledonous plants. Arabidopsis is a genus within the family Brassicaceae, which belongs to the order Capparales within the subclass Dilleniidae. Tobacco,Nicotiana tabacum, belongs to the family Solanaceae, within the order Solanales of the subclass Asteridae. The Dilleniidae are closely related to the Magnoliidae, the most primitive subclass of dicotyledonous plants. In contrast, the Asteridae are the most advanced subclass of dicotyledons (Cronquist, A., An Integrated System of Classification of Flowering Plants, 1981 (New York: Columbia University Press).
- The two familes to which Arabidopsis and tobacco belong, Brassicaceae and Solanaceae, are large familes of major economic importance (Heywood, V.H.,Flowering Plants of the World, 1993, (New York: Oxford University Press). Main crops within the Brassicaceae include oilseed rape and cabbage and its relatives, such as kale, cauliflower, broccoli, and Chinese cabbage. The family Solanaceae is one of the most important serving humankind, containing many essential vegetables and fruits such as potatoes, tomatoes, aubergines, paprika, chilies, and bell peppers.
- Recent work has shown that close homologs of Arabidopsis floral regulatory genes exist in monocotyledonous plants. For example, homologs of the APETALA1 and LEAFY genes have been identified in maize (Veit, et al.,Plant Cell, 5:1205, 1993; Weigel and Meyerowitz, In Molecular Basis of Morphogenesis, pp. 91-105, 1993, (New York: Wiley-Liss).
- Because constitutive expression of LFY can induce flowers precociously during the vegetative phase of Arabidopsis, other species were examined as well. The effect of constitutive LFY expression was studied in a perennial tree, hybrid aspen, which is derived from parental species that flower naturally only after 8-20 years (Schreiner, E.J. inUSDA Agiculture Handbook, 450: Seeds of Woody Plants in the United States (ed. Schopmeyer, C. S.) pp. 645-655 (U.S. Government Printing Office, Washington D.C., 1974). 35S::LFY aspen plants were obtained by Agrobacterium-mediated transformation of stem segments and subsequent regeneration of transgenic shoots in tissue culture (Nilsson, O., et al., Transgen. Res., 1:209, 1992).
- Hybrid aspen was transformed as described previously (Nilsson, O., et al., ibid). Levels of LFY RNA expression were similar to those of 35S::LFY Arabidopsis, as determined by Northern blot analysis. The number of vegetative leaves varied between the different regenerating shoots. Those with a higher number of vegetative leaves formed roots, allowing for transfer to the greenhouse. Individual flowers were removed either from primary transformants that had been transferred to the greenhouse, or from catkins collected in spring 1995 at Carlshem (Umeá, Sweden) from a tree whose age was determined by counting the number of annual rings in a core extracted with an increment borer at 1.5 m above ground level. Flowers were fixed in formaldehyde/acetic acid/ethanol, and destained in ethanol before photography.
- FIG. 6 shows that constitutive expression of Arabidopsis LFY converts aspen shoots into flowers. Panels a and b show five-month-old shoots of hybrid aspen (Populous tremula x tremuloides) grown in tissue culture. Panel a shows a 35S::LFY transformant. Solitary, lateral flowers in the axils of leaves (lf) and an abnormal terminal flower (tf) are indicated. Panel b shows a non-transgenic control. Arrowheads indicate axils of leaves, from which lateral vegetative shoots will emerge, normally in the following year. Note that aspen plants regenerated from tissue culture show the same juvenile phenotype during the first growing cycle as plants grown from seed (Nilsson, O., supra) Panel C is a close-up view of solitary male flower that formed in a leaf axil of a seven-month-old 35S::LFY transformant that had been transferred to the greenhouse. Panel d shows a close-up view of male flower removed from wild-type catkin shown in panel e. Note bract (b) subtending wild-type flower. Panel e shows a cluster of male catkins of P. tremula, one of the parental species of hybrid aspen, taken from a 15-year-old tree. Red pigment in anthers is apparent. Scale bars: a,b, 5 mm; c, d, 1 mm; e, 20 mm.
- Regenerating 35S::LFY aspen shoots initially produced solitary flowers in the axils of normal leaves (FIG. 6a,e). However, the number of vegetative leaves is limited, and the shoot meristem is prematurely consumed in the formation of an aberrant terminal flower (FIG. 6a). Precocious flower development is specific to 35S::LFYtransformants, as such an effect was not observed in non-transgenic controls (FIG. 6b). Furthermore, not a single instance of precocious flower development has been seen in the more than 1,500 other lines of transgenic aspen that were generated with various constructs during the past six years at the Swedish University of Agricultural Sciences (Nilsson O., et al., 1992 supra; Nilsson, 0. Thesis, Swedish University of Agricultural Sciences, 1995).
- Although wild-type Arabidopsis and aspen are rather different, one being a weed and the other a tree, the overall phenotype of 35S::LFY aspen very much resembles that of 35S::LFYArabidopsis. In wild-type plants of both species, flowers are normally formed in lateral positions on inflorescence shoots. In aspen, these inflorescence shoots are called catkins and arise from the leaf axils of adult trees (FIG. 6d, e). In both 35S::LFY Arabidopsis and 35S::LFY aspen, solitary flowers form instead of shoots in the axils of vegetative leaves. Moreover, as in Arabidopsis, the secondary shoots of trangenic aspen are more severely affected than the primary shoot.
- An apparent LFY orthologue from poplar has been described (Strauss, S.H., el al.,Mole. Breed, 1:5, 1995) which, similarly to tobacco LFY (Kelly, A., et al., Pl. Cell, 7:225, 1995) is already expressed during the vegetative stage. The vegetative expression might have suggested that LFY activity is not sufficient to induce flower development in these species. The present results in aspen, which is the same in genus as poplar, indicate that this is not the case, a finding that extends to tobacco, which also flowers very early when transformed with a 35S:: Arabidopsis LFY construct (see Examples 1-3). One possible explanation for the effects of 35S::LFY in these species is that the transgene is expressed at higher effective levels in than the endogenous gene. In tobacco, expression of the endogenous gene in the center of the shoot meristem (which eventually turns into a flower meristem and forms a terminal flower) is relatively low (Kelly, A. J., et al., supra.), and it is conceivable that the other genes, such as AP1, are the primary regulators of flower-meristem-identity in non-transgenic tobacco.
- In addition to LFY, mutations in the genes AP1, CAL, APETALA2 (AP2) and UNUSUAL FLORAL ORGANS (UFO) are known to affect the identity of Arabidopsis flower meristems (Mandel, M.A., et al.,Nature, 360-273, 1992; Irish, V.F. & Sussex, I.M., Pl. Cell, 2:741, 1990; Jofuku, K.D., et al., Pl. Cell, 6, 1994-Levin, J.Z. & Meyerowitz, E.M. Pl. Cell, 7:529, 1995) although lfy mutations have generally the strongest effects, and are the only ones that consistently cause a complete transformation of at least a few flowers into shoots. Mutations in all five genes also affect the identity of floral organs, but meristem-identity and organ-identity defects are at least in some cases separable (Bowman, J.L., et al., Development, 119:721, 1993; Jack, T., et al., Cell, 76:703, 1994). To determine whether any of the other genes are required to mediate the effects of 35S::LFY on meristem identity, or whether their inactivation would merely affect the identity of floral organs in 35S::LFY flowers, the 35S::LFY transgene was crossed into various mutant backgrounds.
- A 35S::LFY tranformant (line DW151.117, Wassilewskija ecotype) was crossed to ap1-1 (Landberg erecta ecotype)(Irish, F.V. & Sussex, I.M., PlCell 2:741, 1990.). Transheterozygote F1 progeny was either backcrossed to ap1-1 or allowed to self-fertilize. The cal genotype of selfed F2 progeny was determined by polymerase chain reaction (PCR)(Kempin, S.A., et al., Science, 267:552, 1994).
- FIG. 7 shows that 35S::LFY phenotype is partly suppressed by an ap1 mutation. Panel a shows five-week-old plants that carry the erecta mutation. The 35S::LFY AP1+plant (left) has no elongated primary shoot. A primary shoot is well developed in the 35S::LFY ap1 plant (middle), although the primary shoot still terminates prematurely, and is shorter than that of the non-transgenic ap1 plant (right). Panels b-d show a detailed view of 35S::LFY ap1 plants. Panel b shows a close-up view of lateral shoot indicated by arrowhead in panel a. Panel c shows emerging shoots in the axils of rosette leaves. Panel d shows a top view of primary shoot with terminal flower (tf). Panels c and d are from a four-week-old plant. The ap1 effects are enhanced further by the cal-1 mutation, although there is no qualitative change in the 35S::LFY ap1 phenotype.
- The ap2-1, ap2-2 and ufo-2 mutations caused only additive phenotypes, and did not significantly affect the shoot-to-flower conversion in 35S::LFY plants. In contrast, the apl-1 mutation suppressed the 35S::LFY phenotype to a notable extent, although terminal flowers were still formed (FIG. 7). Both the primary and secondary shoots were affected, with the strongest effects being observed in lateral positions (FIG. 7b,c). The solitary flowers that develop in the axils of rosette leaves of 35S::LFY AP1+plants become complex shoots with an average of 10 nodes in 35S:LFY apl plants (FIG. 7c). These observations not only confirmed that LFY can induce AP1(which fails to become activated in early arising flowers of lfy mutants), but also that the combined activities of LFY and AP1 are ever more effective in transforming shoot meristems into flower meristems than LFY activity alone.
- Taken together, these results suggest that competence to respond to flower-meristem-identity genes is acquired gradually. In young meristems, competence appears to be low, and both LFY and AP1 are required to promote flower development over that of shoots. Competence increases later in the life cycle, and LFY alone becomes sufficient to induce flower development.
- The present invention shows that constitutive expression of a single flower meristem identity gene, such as LFY or AP1, can induce precocious flower development in plants as diverse as Arabidopsis, an ephermeral weed, and aspen, a perennial tree. The results not only contribute to the understanding of flower development and floral induction, they are also likely to be of interest because shorter flowering times lead to shorter generation times, which in turn allows acceleration of breeding programs. Modern crop varieties are the result of continued improvement by breeding and two recently developed technologies have made breeding even more important. The first technology is molecular mapping, with which genes encoding desirable traits can be rapidly located within the genome. Introduction of such traits into agriculturally important varieties is now greatly assisted by monitoring linked molecular markers, instead of testing for actual expression of these traits. The second technology is transformation of plants with hybrid genes conferring various traits such as engineeered pathogen resistance. However, progress in this area has been delayed because the number of plant varieties amenable to transformation is often restricted, and extensive backcrossing is needed to introgress transgenes into a desired background. In both cases, marker-assisted breeding and transgene introgression, reduction of generation time through the induction of precocious flowering should prove useful.
- The foregoing is meant to illustrate, but not to limit, the scope of the invention. Indeed, those of ordinary skill in the art can readily envision and produce further embodiments, based on the teachings herein, without undue experimentation.
-
1 8 1 1275 DNA Arabdopsis thaliana CDS (1)...(1275) 1 atg gat cct gaa ggt ttc acg agt ggc tta ttc cgg tgg aac cca acg 48 Met Asp Pro Glu Gly Phe Thr Ser Gly Leu Phe Arg Trp Asn Pro Thr 1 5 10 15 aga gca ttg gtt caa gca cca cct ccg gtt cca cct ccg ctg cag caa 96 Arg Ala Leu Val Gln Ala Pro Pro Pro Val Pro Pro Pro Leu Gln Gln 20 25 30 cag ccg gtg aca ccg cag acg gct gct ttt ggg atg cga ctt ggt ggt 144 Gln Pro Val Thr Pro Gln Thr Ala Ala Phe Gly Met Arg Leu Gly Gly 35 40 45 tta gag gga cta ttc ggt cca tac ggt ata cgt ttc tac acg gcg gcg 192 Leu Glu Gly Leu Phe Gly Pro Tyr Gly Ile Arg Phe Tyr Thr Ala Ala 50 55 60 aag ata gcg gag tta ggt ttt acg gcg agc acg ctt gtg ggt atg aag 240 Lys Ile Ala Glu Leu Gly Phe Thr Ala Ser Thr Leu Val Gly Met Lys 65 70 75 80 gac gag gag ctt gaa gag atg atg aat agt ctc tct cat atc ttt cgt 288 Asp Glu Glu Leu Glu Glu Met Met Asn Ser Leu Ser His Ile Phe Arg 85 90 95 tgg gag ctt ctt gtt ggt gaa cgg tac ggt atc aaa gct gcc gtt aga 336 Trp Glu Leu Leu Val Gly Glu Arg Tyr Gly Ile Lys Ala Ala Val Arg 100 105 110 gct gaa cgg aga cga ttg caa gaa gag gag gaa gag gaa tct tct aga 384 Ala Glu Arg Arg Arg Leu Gln Glu Glu Glu Glu Glu Glu Ser Ser Arg 115 120 125 cgc cgt cat ttg cta ctc tcc gcc gct ggt gat tcc ggt act cat cac 432 Arg Arg His Leu Leu Leu Ser Ala Ala Gly Asp Ser Gly Thr His His 130 135 140 gct ctt gat gct ctc tcc caa gaa gat gat tgg aca ggg tta tct gag 480 Ala Leu Asp Ala Leu Ser Gln Glu Asp Asp Trp Thr Gly Leu Ser Glu 145 150 155 160 gaa ccg gtg cag caa caa gac cag act gat gcg gcg ggg aat aac ggc 528 Glu Pro Val Gln Gln Gln Asp Gln Thr Asp Ala Ala Gly Asn Asn Gly 165 170 175 gga gga gga agt ggt tac tgg gac gca ggt caa gga aag atg aag aag 576 Gly Gly Gly Ser Gly Tyr Trp Asp Ala Gly Gln Gly Lys Met Lys Lys 180 185 190 caa cag cag cag aga cgg aga aag aaa cca atg ctg acg tca gtg gaa 624 Gln Gln Gln Gln Arg Arg Arg Lys Lys Pro Met Leu Thr Ser Val Glu 195 200 205 acc gac gaa gac gtc aac gaa ggt gag gat gac gac ggg atg gat aac 672 Thr Asp Glu Asp Val Asn Glu Gly Glu Asp Asp Asp Gly Met Asp Asn 210 215 220 ggc aac gga ggt agt ggt ttg ggg aca gag aga cag agg gag cat ccg 720 Gly Asn Gly Gly Ser Gly Leu Gly Thr Glu Arg Gln Arg Glu His Pro 225 230 235 240 ttt atc gta acg gag cct ggg gaa gtg gca cgt ggc aaa aag aac ggc 768 Phe Ile Val Thr Glu Pro Gly Glu Val Ala Arg Gly Lys Lys Asn Gly 245 250 255 tta gat tat ctg ttc cac ttg tac gaa caa tgc cgt gag ttc ctt ctt 816 Leu Asp Tyr Leu Phe His Leu Tyr Glu Gln Cys Arg Glu Phe Leu Leu 260 265 270 cag gtc cag aca att gct aaa gac cgt ggc gaa aaa tgc ccc acc aag 864 Gln Val Gln Thr Ile Ala Lys Asp Arg Gly Glu Lys Cys Pro Thr Lys 275 280 285 gtg acg aac caa gta ttc agg tac gcg aag aaa tca gga gcg agt tac 912 Val Thr Asn Gln Val Phe Arg Tyr Ala Lys Lys Ser Gly Ala Ser Tyr 290 295 300 ata aac aag cct aaa atg cga cac tac gtt cac tgt tac gct ctc cac 960 Ile Asn Lys Pro Lys Met Arg His Tyr Val His Cys Tyr Ala Leu His 305 310 315 320 tgc cta gac gaa gaa gct tca aat gct ctc aga aga gcg ttt aaa gaa 1008 Cys Leu Asp Glu Glu Ala Ser Asn Ala Leu Arg Arg Ala Phe Lys Glu 325 330 335 cgc ggt gag aac gtt ggc tca tgg cgt cag gct tgt tac aag cca ctt 1056 Arg Gly Glu Asn Val Gly Ser Trp Arg Gln Ala Cys Tyr Lys Pro Leu 340 345 350 gtg aac atc gct tgt cgt cat ggc tgg gat ata gac gcc gtc ttt aac 1104 Val Asn Ile Ala Cys Arg His Gly Trp Asp Ile Asp Ala Val Phe Asn 355 360 365 gct cat cct cgt ctc tct att tgg tat gtt cca aca aag ctg cgt cag 1152 Ala His Pro Arg Leu Ser Ile Trp Tyr Val Pro Thr Lys Leu Arg Gln 370 375 380 ctt tgc cat ttg gag cgg aac aat gcg gtt gct gcg gct gcg gct tta 1200 Leu Cys His Leu Glu Arg Asn Asn Ala Val Ala Ala Ala Ala Ala Leu 385 390 395 400 gtt ggc ggt att agc tgt acc gga tcg tcg acg tct gga cgt ggt gga 1248 Val Gly Gly Ile Ser Cys Thr Gly Ser Ser Thr Ser Gly Arg Gly Gly 405 410 415 tgc ggc ggc gac gac ttg cgt ttc tag 1275 Cys Gly Gly Asp Asp Leu Arg Phe * 420 2 424 PRT Arabdopsis thaliana 2 Met Asp Pro Glu Gly Phe Thr Ser Gly Leu Phe Arg Trp Asn Pro Thr 1 5 10 15 Arg Ala Leu Val Gln Ala Pro Pro Pro Val Pro Pro Pro Leu Gln Gln 20 25 30 Gln Pro Val Thr Pro Gln Thr Ala Ala Phe Gly Met Arg Leu Gly Gly 35 40 45 Leu Glu Gly Leu Phe Gly Pro Tyr Gly Ile Arg Phe Tyr Thr Ala Ala 50 55 60 Lys Ile Ala Glu Leu Gly Phe Thr Ala Ser Thr Leu Val Gly Met Lys 65 70 75 80 Asp Glu Glu Leu Glu Glu Met Met Asn Ser Leu Ser His Ile Phe Arg 85 90 95 Trp Glu Leu Leu Val Gly Glu Arg Tyr Gly Ile Lys Ala Ala Val Arg 100 105 110 Ala Glu Arg Arg Arg Leu Gln Glu Glu Glu Glu Glu Glu Ser Ser Arg 115 120 125 Arg Arg His Leu Leu Leu Ser Ala Ala Gly Asp Ser Gly Thr His His 130 135 140 Ala Leu Asp Ala Leu Ser Gln Glu Asp Asp Trp Thr Gly Leu Ser Glu 145 150 155 160 Glu Pro Val Gln Gln Gln Asp Gln Thr Asp Ala Ala Gly Asn Asn Gly 165 170 175 Gly Gly Gly Ser Gly Tyr Trp Asp Ala Gly Gln Gly Lys Met Lys Lys 180 185 190 Gln Gln Gln Gln Arg Arg Arg Lys Lys Pro Met Leu Thr Ser Val Glu 195 200 205 Thr Asp Glu Asp Val Asn Glu Gly Glu Asp Asp Asp Gly Met Asp Asn 210 215 220 Gly Asn Gly Gly Ser Gly Leu Gly Thr Glu Arg Gln Arg Glu His Pro 225 230 235 240 Phe Ile Val Thr Glu Pro Gly Glu Val Ala Arg Gly Lys Lys Asn Gly 245 250 255 Leu Asp Tyr Leu Phe His Leu Tyr Glu Gln Cys Arg Glu Phe Leu Leu 260 265 270 Gln Val Gln Thr Ile Ala Lys Asp Arg Gly Glu Lys Cys Pro Thr Lys 275 280 285 Val Thr Asn Gln Val Phe Arg Tyr Ala Lys Lys Ser Gly Ala Ser Tyr 290 295 300 Ile Asn Lys Pro Lys Met Arg His Tyr Val His Cys Tyr Ala Leu His 305 310 315 320 Cys Leu Asp Glu Glu Ala Ser Asn Ala Leu Arg Arg Ala Phe Lys Glu 325 330 335 Arg Gly Glu Asn Val Gly Ser Trp Arg Gln Ala Cys Tyr Lys Pro Leu 340 345 350 Val Asn Ile Ala Cys Arg His Gly Trp Asp Ile Asp Ala Val Phe Asn 355 360 365 Ala His Pro Arg Leu Ser Ile Trp Tyr Val Pro Thr Lys Leu Arg Gln 370 375 380 Leu Cys His Leu Glu Arg Asn Asn Ala Val Ala Ala Ala Ala Ala Leu 385 390 395 400 Val Gly Gly Ile Ser Cys Thr Gly Ser Ser Thr Ser Gly Arg Gly Gly 405 410 415 Cys Gly Gly Asp Asp Leu Arg Phe 420 3 1054 DNA Arabdopsis thaliana CDS (124)...(891) 3 ctttccaatt ggttcatacc aaagtctgag ctcttcttta tatctctctt gtagtttctt 60 attgggggtc tttgttttgt ttggttcttt tagagtaaga agtttcttaa aaaaggatca 120 aaa atg gga agg ggt agg gtt caa ttg aag agg ata gag aac aag atc 168 Met Gly Arg Gly Arg Val Gln Leu Lys Arg Ile Glu Asn Lys Ile 1 5 10 15 aat aga caa gtg aca ttc tcg aaa aga aga gct ggt ctt ttg aag aaa 216 Asn Arg Gln Val Thr Phe Ser Lys Arg Arg Ala Gly Leu Leu Lys Lys 20 25 30 gct cat gag atc tct gtt ctc tgt gat gct gaa gtt gct ctt gtt gtc 264 Ala His Glu Ile Ser Val Leu Cys Asp Ala Glu Val Ala Leu Val Val 35 40 45 ttc tcc cat aag ggg aaa ctc ttc gaa tac tcc act gat tct tgt atg 312 Phe Ser His Lys Gly Lys Leu Phe Glu Tyr Ser Thr Asp Ser Cys Met 50 55 60 gag aag ata ctt gaa cgc tat gag agg tac tct tac gcc gaa aga cag 360 Glu Lys Ile Leu Glu Arg Tyr Glu Arg Tyr Ser Tyr Ala Glu Arg Gln 65 70 75 ctt att gca cct gag tcc gac gtc aat aca aac tgg tcg atg gag tat 408 Leu Ile Ala Pro Glu Ser Asp Val Asn Thr Asn Trp Ser Met Glu Tyr 80 85 90 95 aac agg ctt aag gct aag att gag ctt ttg gag aga aac cag agg cat 456 Asn Arg Leu Lys Ala Lys Ile Glu Leu Leu Glu Arg Asn Gln Arg His 100 105 110 tat ctt ggg gaa gac ttg caa gca atg agc cct aaa gag ctt cag aat 504 Tyr Leu Gly Glu Asp Leu Gln Ala Met Ser Pro Lys Glu Leu Gln Asn 115 120 125 ctg gag cag cag ctt gac act gct ctt aag cac atc cgc act aga aaa 552 Leu Glu Gln Gln Leu Asp Thr Ala Leu Lys His Ile Arg Thr Arg Lys 130 135 140 aac caa ctt atg tac gag tcc atc aat gag ctc caa aaa aag gag aag 600 Asn Gln Leu Met Tyr Glu Ser Ile Asn Glu Leu Gln Lys Lys Glu Lys 145 150 155 gcc ata cag gag caa aac agc atg ctt tct aaa cag atc aag gag agg 648 Ala Ile Gln Glu Gln Asn Ser Met Leu Ser Lys Gln Ile Lys Glu Arg 160 165 170 175 gaa aaa att ctt agg gct caa cag gag cag tgg gat cag cag aac caa 696 Glu Lys Ile Leu Arg Ala Gln Gln Glu Gln Trp Asp Gln Gln Asn Gln 180 185 190 ggc cac aat atg cct ccc cct ctg cca ccg cag cag cac caa atc cag 744 Gly His Asn Met Pro Pro Pro Leu Pro Pro Gln Gln His Gln Ile Gln 195 200 205 cat cct tac atg ctc tct cat cag cca tct cct ttt ctc aac atg ggt 792 His Pro Tyr Met Leu Ser His Gln Pro Ser Pro Phe Leu Asn Met Gly 210 215 220 ggt ctg tat caa gaa gat gat cca atg gca atg agg aat gat ctc gaa 840 Gly Leu Tyr Gln Glu Asp Asp Pro Met Ala Met Arg Asn Asp Leu Glu 225 230 235 ctg act ctt gaa ccc gtt tac aac tgc aac ctt ggc tgc ttc gcc gca 888 Leu Thr Leu Glu Pro Val Tyr Asn Cys Asn Leu Gly Cys Phe Ala Ala 240 245 250 255 tga agcatttcca tatatatatt tgtaatcgtc aacaataaaa acagtttgcc 941 * acatacatat aaatagtggc taggctcttt tcatccaatt aatatatttt ggcaaatgtt 1001 cgatgttctt atatcatcat atataaatta gcaggctcct ttcttttttt gta 1054 4 255 PRT Arabdopsis thaliana 4 Met Gly Arg Gly Arg Val Gln Leu Lys Arg Ile Glu Asn Lys Ile Asn 1 5 10 15 Arg Gln Val Thr Phe Ser Lys Arg Arg Ala Gly Leu Leu Lys Lys Ala 20 25 30 His Glu Ile Ser Val Leu Cys Asp Ala Glu Val Ala Leu Val Val Phe 35 40 45 Ser His Lys Gly Lys Leu Phe Glu Tyr Ser Thr Asp Ser Cys Met Glu 50 55 60 Lys Ile Leu Glu Arg Tyr Glu Arg Tyr Ser Tyr Ala Glu Arg Gln Leu 65 70 75 80 Ile Ala Pro Glu Ser Asp Val Asn Thr Asn Trp Ser Met Glu Tyr Asn 85 90 95 Arg Leu Lys Ala Lys Ile Glu Leu Leu Glu Arg Asn Gln Arg His Tyr 100 105 110 Leu Gly Glu Asp Leu Gln Ala Met Ser Pro Lys Glu Leu Gln Asn Leu 115 120 125 Glu Gln Gln Leu Asp Thr Ala Leu Lys His Ile Arg Thr Arg Lys Asn 130 135 140 Gln Leu Met Tyr Glu Ser Ile Asn Glu Leu Gln Lys Lys Glu Lys Ala 145 150 155 160 Ile Gln Glu Gln Asn Ser Met Leu Ser Lys Gln Ile Lys Glu Arg Glu 165 170 175 Lys Ile Leu Arg Ala Gln Gln Glu Gln Trp Asp Gln Gln Asn Gln Gly 180 185 190 His Asn Met Pro Pro Pro Leu Pro Pro Gln Gln His Gln Ile Gln His 195 200 205 Pro Tyr Met Leu Ser His Gln Pro Ser Pro Phe Leu Asn Met Gly Gly 210 215 220 Leu Tyr Gln Glu Asp Asp Pro Met Ala Met Arg Asn Asp Leu Glu Leu 225 230 235 240 Thr Leu Glu Pro Val Tyr Asn Cys Asn Leu Gly Cys Phe Ala Ala 245 250 255 5 1191 DNA Antirihinum majus CDS (1)...(1191) 5 atg gat cct gat gca ttc ttg ttc aaa tgg gac cac aga acc gcc ctc 48 Met Asp Pro Asp Ala Phe Leu Phe Lys Trp Asp His Arg Thr Ala Leu 1 5 10 15 cct caa cca aac agg ctc ctc gac gcc gtg gcc cca ccg cct cct ccg 96 Pro Gln Pro Asn Arg Leu Leu Asp Ala Val Ala Pro Pro Pro Pro Pro 20 25 30 ccg cct cag gcg ccg tca tac tcc atg agg cca aga gaa ctc ggc ggc 144 Pro Pro Gln Ala Pro Ser Tyr Ser Met Arg Pro Arg Glu Leu Gly Gly 35 40 45 tta gaa gaa tta ttc caa gct tat ggc atc aga tac tac act gcc gct 192 Leu Glu Glu Leu Phe Gln Ala Tyr Gly Ile Arg Tyr Tyr Thr Ala Ala 50 55 60 aaa atc gct gaa ctt gga ttc act gtg aac acg ctt ttg gac atg agg 240 Lys Ile Ala Glu Leu Gly Phe Thr Val Asn Thr Leu Leu Asp Met Arg 65 70 75 80 gac gag gag cta gac gag atg atg aac agc ctt tgt cag att ttc agg 288 Asp Glu Glu Leu Asp Glu Met Met Asn Ser Leu Cys Gln Ile Phe Arg 85 90 95 tgg gac cta ctt gtc gga gag agg tat ggg att aag gcg gcg gtg aga 336 Trp Asp Leu Leu Val Gly Glu Arg Tyr Gly Ile Lys Ala Ala Val Arg 100 105 110 gcg gaa cga cgt cgt atc gac gag gag gaa gtg agg cgg agg cat ctc 384 Ala Glu Arg Arg Arg Ile Asp Glu Glu Glu Val Arg Arg Arg His Leu 115 120 125 ttg ttg ggt gat act acg cat gct ctt gat gct ctt tct caa gaa ggg 432 Leu Leu Gly Asp Thr Thr His Ala Leu Asp Ala Leu Ser Gln Glu Gly 130 135 140 ttg tcg gag gag ccg gtg cag caa gaa aag gaa gca atg gga agc ggc 480 Leu Ser Glu Glu Pro Val Gln Gln Glu Lys Glu Ala Met Gly Ser Gly 145 150 155 160 gga ggc ggt gta gga ggc gtg tgg gaa atg atg ggg gcg ggt ggt cga 528 Gly Gly Gly Val Gly Gly Val Trp Glu Met Met Gly Ala Gly Gly Arg 165 170 175 aaa gca ccg cag cgg cgt agg aag aat tac aaa ggg agg tct aga atg 576 Lys Ala Pro Gln Arg Arg Arg Lys Asn Tyr Lys Gly Arg Ser Arg Met 180 185 190 gct tcg atg gag gag gat gat gat gat gat gac gac gaa acc gaa ggg 624 Ala Ser Met Glu Glu Asp Asp Asp Asp Asp Asp Asp Glu Thr Glu Gly 195 200 205 gcg gaa gac gac gaa aat atc gta agc gag cgg cag agg gag cat ccg 672 Ala Glu Asp Asp Glu Asn Ile Val Ser Glu Arg Gln Arg Glu His Pro 210 215 220 ttt atc gtg acg gag ccc gga gag gtg gcg cgt ggg aaa aag aat ggt 720 Phe Ile Val Thr Glu Pro Gly Glu Val Ala Arg Gly Lys Lys Asn Gly 225 230 235 240 ctt gat tat ttg ttt cat ttg tac gag caa tgc cgc gac ttc ttg atc 768 Leu Asp Tyr Leu Phe His Leu Tyr Glu Gln Cys Arg Asp Phe Leu Ile 245 250 255 caa gtt caa act att gct aag gag aga ggt gaa aaa tgt ccc act aag 816 Gln Val Gln Thr Ile Ala Lys Glu Arg Gly Glu Lys Cys Pro Thr Lys 260 265 270 gtg acg aac caa gtg ttc agg tac gca aag aag gct ggc gct aac tac 864 Val Thr Asn Gln Val Phe Arg Tyr Ala Lys Lys Ala Gly Ala Asn Tyr 275 280 285 atc aac aaa cca aaa atg cgc cac tac gtg cac tgc tac gcc ctg cac 912 Ile Asn Lys Pro Lys Met Arg His Tyr Val His Cys Tyr Ala Leu His 290 295 300 tgc ctt gat gag gcc gcg tcc aat gca ctt cgt cgg gca ttc aag gag 960 Cys Leu Asp Glu Ala Ala Ser Asn Ala Leu Arg Arg Ala Phe Lys Glu 305 310 315 320 cgt ggt gag aac gtc ggt gca tgg cgt cag gca tgc tac aag ccc ttg 1008 Arg Gly Glu Asn Val Gly Ala Trp Arg Gln Ala Cys Tyr Lys Pro Leu 325 330 335 gtg gcc att gca gca aga caa gga tgg gat atc gat acc ata ttc aac 1056 Val Ala Ile Ala Ala Arg Gln Gly Trp Asp Ile Asp Thr Ile Phe Asn 340 345 350 gct cat ccc cgt ctc tcg atc tgg tat gtc ccc acc aag ctt cgt cag 1104 Ala His Pro Arg Leu Ser Ile Trp Tyr Val Pro Thr Lys Leu Arg Gln 355 360 365 ctc tgc cat gcc gag agg agc agt gcg gca gtt gct gcc acc agc tcc 1152 Leu Cys His Ala Glu Arg Ser Ser Ala Ala Val Ala Ala Thr Ser Ser 370 375 380 atc acc gga ggt ggg ccg gca gat cac ttg ccg ttt tag 1191 Ile Thr Gly Gly Gly Pro Ala Asp His Leu Pro Phe * 385 390 395 6 396 PRT Antirihinum majus 6 Met Asp Pro Asp Ala Phe Leu Phe Lys Trp Asp His Arg Thr Ala Leu 1 5 10 15 Pro Gln Pro Asn Arg Leu Leu Asp Ala Val Ala Pro Pro Pro Pro Pro 20 25 30 Pro Pro Gln Ala Pro Ser Tyr Ser Met Arg Pro Arg Glu Leu Gly Gly 35 40 45 Leu Glu Glu Leu Phe Gln Ala Tyr Gly Ile Arg Tyr Tyr Thr Ala Ala 50 55 60 Lys Ile Ala Glu Leu Gly Phe Thr Val Asn Thr Leu Leu Asp Met Arg 65 70 75 80 Asp Glu Glu Leu Asp Glu Met Met Asn Ser Leu Cys Gln Ile Phe Arg 85 90 95 Trp Asp Leu Leu Val Gly Glu Arg Tyr Gly Ile Lys Ala Ala Val Arg 100 105 110 Ala Glu Arg Arg Arg Ile Asp Glu Glu Glu Val Arg Arg Arg His Leu 115 120 125 Leu Leu Gly Asp Thr Thr His Ala Leu Asp Ala Leu Ser Gln Glu Gly 130 135 140 Leu Ser Glu Glu Pro Val Gln Gln Glu Lys Glu Ala Met Gly Ser Gly 145 150 155 160 Gly Gly Gly Val Gly Gly Val Trp Glu Met Met Gly Ala Gly Gly Arg 165 170 175 Lys Ala Pro Gln Arg Arg Arg Lys Asn Tyr Lys Gly Arg Ser Arg Met 180 185 190 Ala Ser Met Glu Glu Asp Asp Asp Asp Asp Asp Asp Glu Thr Glu Gly 195 200 205 Ala Glu Asp Asp Glu Asn Ile Val Ser Glu Arg Gln Arg Glu His Pro 210 215 220 Phe Ile Val Thr Glu Pro Gly Glu Val Ala Arg Gly Lys Lys Asn Gly 225 230 235 240 Leu Asp Tyr Leu Phe His Leu Tyr Glu Gln Cys Arg Asp Phe Leu Ile 245 250 255 Gln Val Gln Thr Ile Ala Lys Glu Arg Gly Glu Lys Cys Pro Thr Lys 260 265 270 Val Thr Asn Gln Val Phe Arg Tyr Ala Lys Lys Ala Gly Ala Asn Tyr 275 280 285 Ile Asn Lys Pro Lys Met Arg His Tyr Val His Cys Tyr Ala Leu His 290 295 300 Cys Leu Asp Glu Ala Ala Ser Asn Ala Leu Arg Arg Ala Phe Lys Glu 305 310 315 320 Arg Gly Glu Asn Val Gly Ala Trp Arg Gln Ala Cys Tyr Lys Pro Leu 325 330 335 Val Ala Ile Ala Ala Arg Gln Gly Trp Asp Ile Asp Thr Ile Phe Asn 340 345 350 Ala His Pro Arg Leu Ser Ile Trp Tyr Val Pro Thr Lys Leu Arg Gln 355 360 365 Leu Cys His Ala Glu Arg Ser Ser Ala Ala Val Ala Ala Thr Ser Ser 370 375 380 Ile Thr Gly Gly Gly Pro Ala Asp His Leu Pro Phe 385 390 395 7 747 DNA Antirihinum majus CDS (1)...(747) 7 atg ggg aga ggg aaa gta caa ctg aag agg ata gag aac aag atc aat 48 Met Gly Arg Gly Lys Val Gln Leu Lys Arg Ile Glu Asn Lys Ile Asn 1 5 10 15 aga cag gtg act ttc tca aag agg aga ggt cca ttg ttg aaa aaa gct 96 Arg Gln Val Thr Phe Ser Lys Arg Arg Gly Pro Leu Leu Lys Lys Ala 20 25 30 cat gag ctc tct gtg ctt tgt gat gct gaa gtg gct ctt att gtc ttc 144 His Glu Leu Ser Val Leu Cys Asp Ala Glu Val Ala Leu Ile Val Phe 35 40 45 tct aat aag ggg aag cta ttt gag tat tct act gat tct tgc atg gac 192 Ser Asn Lys Gly Lys Leu Phe Glu Tyr Ser Thr Asp Ser Cys Met Asp 50 55 60 agg atc ctg gag aag tat gaa agg tat tca ttt gca gaa aga cag tta 240 Arg Ile Leu Glu Lys Tyr Glu Arg Tyr Ser Phe Ala Glu Arg Gln Leu 65 70 75 80 gtt tca aat gaa cct cag tca cct gcg aat tgg acc ctc gaa tac agc 288 Val Ser Asn Glu Pro Gln Ser Pro Ala Asn Trp Thr Leu Glu Tyr Ser 85 90 95 aaa ctg aag gca aga att gag ctc ttg caa aga aac cat agg cac tat 336 Lys Leu Lys Ala Arg Ile Glu Leu Leu Gln Arg Asn His Arg His Tyr 100 105 110 atg gga gaa gat ctg gac tcc atg agc ctc aaa gag att cag agt cta 384 Met Gly Glu Asp Leu Asp Ser Met Ser Leu Lys Glu Ile Gln Ser Leu 115 120 125 gaa caa cag ctg gac act gct ctt aag aac att cgg acc aga aaa aac 432 Glu Gln Gln Leu Asp Thr Ala Leu Lys Asn Ile Arg Thr Arg Lys Asn 130 135 140 cag ctc ttg tac gat tca atc tct gaa ttg cag cat aag gag aag gca 480 Gln Leu Leu Tyr Asp Ser Ile Ser Glu Leu Gln His Lys Glu Lys Ala 145 150 155 160 ata caa gag caa aac acc atg ctg gca aag aag atc aaa gag aag gag 528 Ile Gln Glu Gln Asn Thr Met Leu Ala Lys Lys Ile Lys Glu Lys Glu 165 170 175 aag gaa att gca caa cag cca cag tgg gag cat cat cgc cac cac act 576 Lys Glu Ile Ala Gln Gln Pro Gln Trp Glu His His Arg His His Thr 180 185 190 aat gca tcg att atg cca ccg cca cca caa tat tcc atg gca cct caa 624 Asn Ala Ser Ile Met Pro Pro Pro Pro Gln Tyr Ser Met Ala Pro Gln 195 200 205 ttc ccc tgc ata aat gtc gga aac aca tat gaa gga gaa gga gca aat 672 Phe Pro Cys Ile Asn Val Gly Asn Thr Tyr Glu Gly Glu Gly Ala Asn 210 215 220 gag gat aga aga aat gag ctt gac ctc act ctt gat tca ctc tat tca 720 Glu Asp Arg Arg Asn Glu Leu Asp Leu Thr Leu Asp Ser Leu Tyr Ser 225 230 235 240 tgc cat ctt gga tgc ttt gct gca tga 747 Cys His Leu Gly Cys Phe Ala Ala * 245 8 248 PRT Antirihinum majus 8 Met Gly Arg Gly Lys Val Gln Leu Lys Arg Ile Glu Asn Lys Ile Asn 1 5 10 15 Arg Gln Val Thr Phe Ser Lys Arg Arg Gly Pro Leu Leu Lys Lys Ala 20 25 30 His Glu Leu Ser Val Leu Cys Asp Ala Glu Val Ala Leu Ile Val Phe 35 40 45 Ser Asn Lys Gly Lys Leu Phe Glu Tyr Ser Thr Asp Ser Cys Met Asp 50 55 60 Arg Ile Leu Glu Lys Tyr Glu Arg Tyr Ser Phe Ala Glu Arg Gln Leu 65 70 75 80 Val Ser Asn Glu Pro Gln Ser Pro Ala Asn Trp Thr Leu Glu Tyr Ser 85 90 95 Lys Leu Lys Ala Arg Ile Glu Leu Leu Gln Arg Asn His Arg His Tyr 100 105 110 Met Gly Glu Asp Leu Asp Ser Met Ser Leu Lys Glu Ile Gln Ser Leu 115 120 125 Glu Gln Gln Leu Asp Thr Ala Leu Lys Asn Ile Arg Thr Arg Lys Asn 130 135 140 Gln Leu Leu Tyr Asp Ser Ile Ser Glu Leu Gln His Lys Glu Lys Ala 145 150 155 160 Ile Gln Glu Gln Asn Thr Met Leu Ala Lys Lys Ile Lys Glu Lys Glu 165 170 175 Lys Glu Ile Ala Gln Gln Pro Gln Trp Glu His His Arg His His Thr 180 185 190 Asn Ala Ser Ile Met Pro Pro Pro Pro Gln Tyr Ser Met Ala Pro Gln 195 200 205 Phe Pro Cys Ile Asn Val Gly Asn Thr Tyr Glu Gly Glu Gly Ala Asn 210 215 220 Glu Asp Arg Arg Asn Glu Leu Asp Leu Thr Leu Asp Ser Leu Tyr Ser 225 230 235 240 Cys His Leu Gly Cys Phe Ala Ala 245
Claims (21)
Priority Applications (1)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
US09/849,772 US20030167539A1 (en) | 1994-12-21 | 2001-05-04 | Genetically modified plants having modulated flower development |
Applications Claiming Priority (4)
Application Number | Priority Date | Filing Date | Title |
---|---|---|---|
US08/360,336 US5637785A (en) | 1994-12-21 | 1994-12-21 | Genetically modified plants having modulated flower development |
US08/576,156 US5844119A (en) | 1994-12-21 | 1995-12-21 | Genetically modified plants having modulated flower development |
US20409498A | 1998-12-01 | 1998-12-01 | |
US09/849,772 US20030167539A1 (en) | 1994-12-21 | 2001-05-04 | Genetically modified plants having modulated flower development |
Related Parent Applications (1)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
US20409498A Continuation | 1994-12-21 | 1998-12-01 |
Publications (1)
Publication Number | Publication Date |
---|---|
US20030167539A1 true US20030167539A1 (en) | 2003-09-04 |
Family
ID=23417554
Family Applications (3)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
US08/360,336 Expired - Lifetime US5637785A (en) | 1994-12-21 | 1994-12-21 | Genetically modified plants having modulated flower development |
US08/576,156 Expired - Fee Related US5844119A (en) | 1994-12-21 | 1995-12-21 | Genetically modified plants having modulated flower development |
US09/849,772 Abandoned US20030167539A1 (en) | 1994-12-21 | 2001-05-04 | Genetically modified plants having modulated flower development |
Family Applications Before (2)
Application Number | Title | Priority Date | Filing Date |
---|---|---|---|
US08/360,336 Expired - Lifetime US5637785A (en) | 1994-12-21 | 1994-12-21 | Genetically modified plants having modulated flower development |
US08/576,156 Expired - Fee Related US5844119A (en) | 1994-12-21 | 1995-12-21 | Genetically modified plants having modulated flower development |
Country Status (10)
Country | Link |
---|---|
US (3) | US5637785A (en) |
EP (1) | EP0798958A4 (en) |
JP (1) | JPH10513042A (en) |
AU (1) | AU711551B2 (en) |
BR (1) | BR9510172A (en) |
CA (1) | CA2208061A1 (en) |
FI (1) | FI972581A (en) |
NO (1) | NO972903L (en) |
NZ (1) | NZ301454A (en) |
WO (1) | WO1996019105A1 (en) |
Families Citing this family (129)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
WO1997025433A1 (en) * | 1996-01-09 | 1997-07-17 | Eidg. Technische Hochschule Zürich Ethz | Regulation of flowering in plants |
US6025543A (en) * | 1996-06-05 | 2000-02-15 | The Regents Of The University Of California | Seed plants exhibiting early reproductive development and methods of making same |
US6002069A (en) * | 1996-06-05 | 1999-12-14 | The Regents Of The University Of California | Seed plants exhibiting inducible early reproductive development and methods of making same |
US6025483A (en) * | 1996-06-05 | 2000-02-15 | The Regents Of The University Of California | Maize and cauliflower apetalai gene products and nucleic acid molecules encoding same |
US6395892B1 (en) | 1998-04-06 | 2002-05-28 | The State Of Oregon Acting By And Through The State Board Of Higher Education On Behalf Of Oregon State University | Floral homeotic genes for manipulation of flowering in poplar and other plant species |
US6693228B1 (en) | 1999-02-25 | 2004-02-17 | Wisconsin Alumni Research Foundation | Alteration of flowering time in plants |
US6828478B2 (en) | 2001-05-09 | 2004-12-07 | The Regents Of The University Of California | Combinations of genes for producing seed plants exhibiting modulated reproductive development |
AU2002364613A1 (en) * | 2001-12-31 | 2003-07-24 | Yeda Research And Development Co., Ltd | A method to maintain a genic male-sterile female parental line of wheat through selfing of the maintainer line |
EP1781083A4 (en) * | 2004-06-18 | 2009-09-09 | Univ California | Brassica indehiscent1 sequences |
DK1763582T3 (en) | 2004-07-08 | 2015-01-12 | Dlf Trifolium As | Means and method of controlling the flowering of plants |
WO2008121291A1 (en) | 2007-03-28 | 2008-10-09 | Monsanto Technology Llc | Utility of snp markers associated with major soybean plant maturity and growth habit genomic regions |
WO2009035852A2 (en) | 2007-09-11 | 2009-03-19 | Monsanto Technology Llc | Increased alpha-prime beta-conglycinin soybeans |
WO2009102890A2 (en) * | 2008-02-12 | 2009-08-20 | Dow Agrosciences Llc | Dominant earliness mutation and gene in sunflower (helianthus annuus) |
US8955523B2 (en) | 2010-01-15 | 2015-02-17 | R.J. Reynolds Tobacco Company | Tobacco-derived components and materials |
US9039839B2 (en) | 2010-04-08 | 2015-05-26 | R.J. Reynolds Tobacco Company | Smokeless tobacco composition comprising tobacco-derived material and non-tobacco plant material |
US9402415B2 (en) | 2010-04-21 | 2016-08-02 | R. J. Reynolds Tobacco Company | Tobacco seed-derived components and materials |
US11116237B2 (en) | 2010-08-11 | 2021-09-14 | R.J. Reynolds Tobacco Company | Meltable smokeless tobacco composition |
US9155321B2 (en) | 2010-08-11 | 2015-10-13 | R.J. Reynolds Tobacco Company | Meltable smokeless tobacco composition |
US9675102B2 (en) | 2010-09-07 | 2017-06-13 | R. J. Reynolds Tobacco Company | Smokeless tobacco product comprising effervescent composition |
US20120125354A1 (en) | 2010-11-18 | 2012-05-24 | R.J. Reynolds Tobacco Company | Fire-Cured Tobacco Extract and Tobacco Products Made Therefrom |
US9220295B2 (en) | 2010-12-01 | 2015-12-29 | R.J. Reynolds Tobacco Company | Tobacco separation process for extracting tobacco-derived materials, and associated extraction systems |
US9204667B2 (en) | 2010-12-01 | 2015-12-08 | R.J. Reynolds Tobacco Company | Smokeless tobacco pastille and injection molding process for forming smokeless tobacco products |
US9775376B2 (en) | 2010-12-01 | 2017-10-03 | R.J. Reynolds Tobacco Company | Smokeless tobacco pastille and moulding process for forming smokeless tobacco products |
US20120152265A1 (en) | 2010-12-17 | 2012-06-21 | R.J. Reynolds Tobacco Company | Tobacco-Derived Syrup Composition |
US8893725B2 (en) | 2011-01-28 | 2014-11-25 | R. J. Reynolds Tobacco Company | Polymeric materials derived from tobacco |
US9107453B2 (en) | 2011-01-28 | 2015-08-18 | R.J. Reynolds Tobacco Company | Tobacco-derived casing composition |
US9254001B2 (en) | 2011-04-27 | 2016-02-09 | R.J. Reynolds Tobacco Company | Tobacco-derived components and materials |
US9192193B2 (en) | 2011-05-19 | 2015-11-24 | R.J. Reynolds Tobacco Company | Molecularly imprinted polymers for treating tobacco material and filtering smoke from smoking articles |
US9474303B2 (en) | 2011-09-22 | 2016-10-25 | R.J. Reynolds Tobacco Company | Translucent smokeless tobacco product |
US20130118512A1 (en) | 2011-11-16 | 2013-05-16 | R.J. Reynolds Tobacco Company | Smokeless tobacco products with starch component |
US20130125907A1 (en) | 2011-11-17 | 2013-05-23 | Michael Francis Dube | Method for Producing Triethyl Citrate from Tobacco |
US20130125904A1 (en) | 2011-11-18 | 2013-05-23 | R.J. Reynolds Tobacco Company | Smokeless tobacco product comprising pectin component |
US10881132B2 (en) | 2011-12-14 | 2021-01-05 | R.J. Reynolds Tobacco Company | Smokeless tobacco product comprising effervescent composition |
US20130206150A1 (en) | 2012-02-10 | 2013-08-15 | R.J. Reynolds Tobacco Company | Multi-layer smokeless tobacco composition |
US9420825B2 (en) | 2012-02-13 | 2016-08-23 | R.J. Reynolds Tobacco Company | Whitened tobacco composition |
EP2827727B1 (en) | 2012-03-19 | 2021-10-13 | R. J. Reynolds Tobacco Company | Method for treating an extracted tobacco pulp and tobacco products made therefrom |
US20130269719A1 (en) | 2012-04-11 | 2013-10-17 | R.J. Reynolds Tobacco Company | Method for treating plants with probiotics |
US9485953B2 (en) | 2012-07-19 | 2016-11-08 | R.J. Reynolds Tobacco Company | Method for treating tobacco plants with enzymes |
US9289011B2 (en) | 2013-03-07 | 2016-03-22 | R.J. Reynolds Tobacco Company | Method for producing lutein from tobacco |
US9155334B2 (en) | 2013-04-05 | 2015-10-13 | R.J. Reynolds Tobacco Company | Modification of bacterial profile of tobacco |
US20150034109A1 (en) | 2013-08-02 | 2015-02-05 | R.J. Reynolds Tobacco Company | Process for Producing Lignin from Tobacco |
US10357054B2 (en) | 2013-10-16 | 2019-07-23 | R.J. Reynolds Tobacco Company | Smokeless tobacco pastille |
US9265284B2 (en) | 2014-01-17 | 2016-02-23 | R.J. Reynolds Tobacco Company | Process for producing flavorants and related materials |
US20160073686A1 (en) | 2014-09-12 | 2016-03-17 | R.J. Reynolds Tobacco Company | Tobacco-derived filter element |
US11219244B2 (en) | 2014-12-22 | 2022-01-11 | R.J. Reynolds Tobacco Company | Tobacco-derived carbon material |
US10881133B2 (en) | 2015-04-16 | 2021-01-05 | R.J. Reynolds Tobacco Company | Tobacco-derived cellulosic sugar |
US20170055565A1 (en) | 2015-09-02 | 2017-03-02 | R. J. Reynolds Tobacco Company | Systems and Apparatus for Reducing Tobacco-Specific Nitrosamines in Dark-Fire Cured Tobacco Through Electronic Control of Curing Conditions |
US20170059554A1 (en) | 2015-09-02 | 2017-03-02 | R. J. Reynolds Tobacco Company | Method for monitoring use of a tobacco product |
US10499684B2 (en) | 2016-01-28 | 2019-12-10 | R.J. Reynolds Tobacco Company | Tobacco-derived flavorants |
US11154087B2 (en) | 2016-02-02 | 2021-10-26 | R.J. Reynolds Tobacco Company | Method for preparing flavorful compounds isolated from black liquor and products incorporating the flavorful compounds |
US10721957B2 (en) | 2016-10-04 | 2020-07-28 | R.J. Reynolds Tobacco Company | Tobacco-derived colorants and colored substrates |
US10813383B2 (en) | 2016-12-12 | 2020-10-27 | R.J. Reynolds Tobacco Company | Dehydration of tobacco and tobacco-derived materials |
US10196778B2 (en) | 2017-03-20 | 2019-02-05 | R.J. Reynolds Tobacco Company | Tobacco-derived nanocellulose material |
US10470487B2 (en) | 2017-04-06 | 2019-11-12 | R.J. Reynolds Tobacco Company | Smoke treatment |
JP2020529206A (en) | 2017-07-31 | 2020-10-08 | アール・ジエイ・レイノルズ・タバコ・カンパニー | Methods and compositions for virus-based gene editing in plants |
BR112020004764A2 (en) | 2017-09-11 | 2020-09-24 | R.J. Reynolds Tobacco Company | methods and compositions to increase the expression of genes of interest in a plant by coexpression with p21 |
US11278050B2 (en) | 2017-10-20 | 2022-03-22 | R.J. Reynolds Tobacco Company | Methods for treating tobacco and tobacco-derived materials to reduce nitrosamines |
WO2019239356A1 (en) | 2018-06-15 | 2019-12-19 | R. J. Reynolds Tobacco Company | Purification of nicotine |
US20200196658A1 (en) | 2018-12-20 | 2020-06-25 | R.J. Reynolds Tobacco Company | Method for whitening tobacco |
CN110343704B (en) * | 2019-05-08 | 2020-12-11 | 中国科学院植物研究所 | AP1 gene mutant and method for regulating and controlling plant calyx and petal opening time |
US11213062B2 (en) | 2019-05-09 | 2022-01-04 | American Snuff Company | Stabilizer for moist snuff |
US20210068446A1 (en) | 2019-09-11 | 2021-03-11 | R. J. Reynolds Tobacco Company | Oral product with cellulosic flavor stabilizer |
WO2021048770A1 (en) | 2019-09-11 | 2021-03-18 | Nicoventures Trading Limited | Alternative methods for whitening tobacco |
US20210068448A1 (en) | 2019-09-11 | 2021-03-11 | Nicoventures Trading Limited | Method for whitening tobacco |
US20210068447A1 (en) | 2019-09-11 | 2021-03-11 | R. J. Reynolds Tobacco Company | Pouched products with enhanced flavor stability |
US11369131B2 (en) | 2019-09-13 | 2022-06-28 | Nicoventures Trading Limited | Method for whitening tobacco |
US11903406B2 (en) | 2019-09-18 | 2024-02-20 | American Snuff Company, Llc | Method for fermenting tobacco |
WO2021086367A1 (en) | 2019-10-31 | 2021-05-06 | Nicoventures Trading Limited | Oral product and method of manufacture |
US20210169126A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral composition with salt inclusion |
US20210169137A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Pouched products |
WO2021116842A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Oral products with controlled release |
WO2021116881A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Oral product in a pourous pouch comprising a fleece material |
MX2022006980A (en) | 2019-12-09 | 2022-08-25 | Nicoventures Trading Ltd | Pouched products with heat sealable binder. |
WO2021116916A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Oral product with multiple flavors having different release profiles |
WO2021116855A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Oral compositions and methods of manufacture |
US20210169868A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral compositions with reduced water content |
US20210169121A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Liquid oral composition |
US20210169784A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Buffered oral compositions |
US20210169132A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral composition including gels |
US20210169129A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Lipid-containing oral composition |
US20210169890A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral composition with polymeric component |
WO2021116895A2 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Stimulus-responsive pouch |
US11889856B2 (en) | 2019-12-09 | 2024-02-06 | Nicoventures Trading Limited | Oral foam composition |
US20210169783A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral products with controlled release |
US20210169788A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral product and method of manufacture |
US11793230B2 (en) | 2019-12-09 | 2023-10-24 | Nicoventures Trading Limited | Oral products with improved binding of active ingredients |
US20210169138A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Fibrous fleece material |
US11672862B2 (en) | 2019-12-09 | 2023-06-13 | Nicoventures Trading Limited | Oral products with reduced irritation |
WO2021116865A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Agents for oral composition |
US20210169123A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Pouched products with enhanced flavor stability |
WO2021116919A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Fleece for oral product with releasable component |
US20210170031A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral composition with nanocrystalline cellulose |
US11617744B2 (en) | 2019-12-09 | 2023-04-04 | Nico Ventures Trading Limited | Moist oral compositions |
US20210169786A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral composition with beet material |
US20210169785A1 (en) | 2019-12-09 | 2021-06-10 | Nicoventures Trading Limited | Oral compositions with reduced water activity |
WO2021116856A2 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Oral products |
WO2021116884A1 (en) | 2019-12-09 | 2021-06-17 | Nicoventures Trading Limited | Layered fleece for pouched product |
EP4072334A1 (en) | 2019-12-09 | 2022-10-19 | Nicoventures Trading Limited | Oral product with dissolvable component |
MX2022007085A (en) | 2019-12-09 | 2022-08-19 | Nicoventures Trading Ltd | Nanoemulsion for oral use. |
US11712059B2 (en) | 2020-02-24 | 2023-08-01 | Nicoventures Trading Limited | Beaded tobacco material and related method of manufacture |
WO2021250516A1 (en) | 2020-06-08 | 2021-12-16 | Nicoventures Trading Limited | Effervescent oral composition comprising an active ingredient |
US11937626B2 (en) | 2020-09-04 | 2024-03-26 | Nicoventures Trading Limited | Method for whitening tobacco |
WO2022053982A1 (en) | 2020-09-11 | 2022-03-17 | Nicoventures Trading Limited | Alginate-based substrates |
US20240008522A1 (en) | 2020-11-18 | 2024-01-11 | Nicoventures Trading Limited | Oral products |
EP4284972A1 (en) | 2021-01-28 | 2023-12-06 | Nicoventures Trading Limited | Method for sealing pouches |
JP2024510316A (en) | 2021-03-19 | 2024-03-06 | ニコベンチャーズ トレーディング リミテッド | Extruded substrates for aerosol delivery devices |
CA3212627A1 (en) | 2021-03-19 | 2022-09-22 | Caroline W. CLARK | Beaded substrates for aerosol delivery devices |
CA3216322A1 (en) | 2021-04-22 | 2022-10-27 | Nicoventures Trading Limited | Effervescent oral composition |
JP2024515358A (en) | 2021-04-22 | 2024-04-09 | ニコベンチャーズ トレーディング リミテッド | Oral cavity composition and manufacturing method |
US20220346436A1 (en) | 2021-04-22 | 2022-11-03 | Nicoventures Trading Limited | Orally dissolving films |
US20220354785A1 (en) | 2021-04-22 | 2022-11-10 | Nicoventures Trading Limited | Oral lozenge products |
US20220354155A1 (en) | 2021-04-30 | 2022-11-10 | Nicoventures Trading Limited | Multi-compartment oral pouched product |
WO2022229929A1 (en) | 2021-04-30 | 2022-11-03 | Nicoventures Trading Limited | Oral products with high-density load |
US20220369688A1 (en) | 2021-05-06 | 2022-11-24 | Nicoventures Trading Limited | Oral compositions and related methods for reducing throat irritation |
CA3222813A1 (en) | 2021-06-16 | 2022-12-22 | Anthony Richard Gerardi | Pouched product comprising dissolvable composition |
CA3223460A1 (en) | 2021-06-21 | 2022-12-29 | Nicoventures Trading Limited | Oral product tablet and method of manufacture |
WO2022269556A1 (en) | 2021-06-25 | 2022-12-29 | Nicoventures Trading Limited | Oral products and method of manufacture |
AU2022306261A1 (en) | 2021-07-09 | 2024-02-29 | Nicoventures Trading Limited | Extruded structures |
WO2023007440A1 (en) | 2021-07-30 | 2023-02-02 | Nicoventures Trading Limited | Aerosol generating substrate comprising microcrystalline cellulose |
US20230098503A1 (en) | 2021-09-30 | 2023-03-30 | Nicoventures Trading Limited | Oral gum composition |
WO2023053062A1 (en) | 2021-09-30 | 2023-04-06 | Nicoventures Trading Limited | Oral product with a basic amine and an ion pairing agent |
WO2023084499A1 (en) | 2021-11-15 | 2023-05-19 | Nicoventures Trading Limited | Products with enhanced sensory characteristics |
US20230148652A1 (en) | 2021-11-15 | 2023-05-18 | Nicoventures Trading Limited | Oral products with nicotine-polymer complex |
WO2023119134A1 (en) | 2021-12-20 | 2023-06-29 | Nicoventures Trading Limited | Substrate material comprising beads for aerosol delivery devices |
US20230309603A1 (en) | 2022-03-31 | 2023-10-05 | R.J. Reynolds Tobacco Company | Agglomerated botanical material for oral products |
WO2023194959A1 (en) | 2022-04-06 | 2023-10-12 | Nicoventures Trading Limited | Pouched products with heat sealable binder |
WO2023242822A1 (en) | 2022-06-17 | 2023-12-21 | Nicoventures Trading Limited | Tobacco-coated sheet and consumable made therefrom |
WO2024069542A1 (en) | 2022-09-30 | 2024-04-04 | R. J. Reynolds Tobacco Company | Method for forming reconstituted tobacco |
WO2024069544A1 (en) | 2022-09-30 | 2024-04-04 | Nicoventures Trading Limited | Reconstituted tobacco substrate for aerosol delivery device |
Family Cites Families (2)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
US4513532A (en) * | 1983-04-14 | 1985-04-30 | Cornnuts Hybrids, Inc. | Genetic factor capable of altering leaf number and distribution in maize |
EP0672155A1 (en) * | 1992-06-30 | 1995-09-20 | Asgrow Seed Company | A method for obtaining a plant having altered floral morphology and a method for protecting plants against pest insects |
-
1994
- 1994-12-21 US US08/360,336 patent/US5637785A/en not_active Expired - Lifetime
-
1995
- 1995-12-21 CA CA002208061A patent/CA2208061A1/en not_active Abandoned
- 1995-12-21 BR BR9510172A patent/BR9510172A/en not_active IP Right Cessation
- 1995-12-21 WO PCT/US1995/016859 patent/WO1996019105A1/en not_active Application Discontinuation
- 1995-12-21 EP EP95944528A patent/EP0798958A4/en not_active Withdrawn
- 1995-12-21 JP JP8520030A patent/JPH10513042A/en active Pending
- 1995-12-21 US US08/576,156 patent/US5844119A/en not_active Expired - Fee Related
- 1995-12-21 NZ NZ301454A patent/NZ301454A/en unknown
- 1995-12-21 AU AU46884/96A patent/AU711551B2/en not_active Ceased
-
1997
- 1997-06-17 FI FI972581A patent/FI972581A/en unknown
- 1997-06-20 NO NO972903A patent/NO972903L/en not_active Application Discontinuation
-
2001
- 2001-05-04 US US09/849,772 patent/US20030167539A1/en not_active Abandoned
Also Published As
Publication number | Publication date |
---|---|
BR9510172A (en) | 1997-10-14 |
JPH10513042A (en) | 1998-12-15 |
EP0798958A1 (en) | 1997-10-08 |
AU4688496A (en) | 1996-07-10 |
US5844119A (en) | 1998-12-01 |
WO1996019105A1 (en) | 1996-06-27 |
AU711551B2 (en) | 1999-10-14 |
CA2208061A1 (en) | 1996-06-27 |
FI972581A (en) | 1997-08-19 |
EP0798958A4 (en) | 1998-05-27 |
NZ301454A (en) | 1999-08-30 |
NO972903D0 (en) | 1997-06-20 |
FI972581A0 (en) | 1997-06-17 |
NO972903L (en) | 1997-08-18 |
US5637785A (en) | 1997-06-10 |
Similar Documents
Publication | Publication Date | Title |
---|---|---|
US5844119A (en) | Genetically modified plants having modulated flower development | |
US7692067B2 (en) | Yield and stress tolerance in transgenic plants | |
US6248937B1 (en) | Transcription factor and method for regulation of seed development, quality and stress-tolerance | |
US5880330A (en) | Shoot meristem specific promoter sequences | |
AU757050B2 (en) | Apomixis conferred by expression of SERK interacting proteins | |
Uberlacker et al. | Ectopic expression of the maize homeobox genes ZmHox1a or ZmHox1b causes pleiotropic alterations in the vegetative and floral development of transgenic tobacco. | |
AU2006314535A1 (en) | EMP4 gene | |
US20100138962A1 (en) | Use of plant chromatin remodeling genes for modulating plant architecture and growth | |
US6630616B1 (en) | Arabidopsis MPC1 gene and methods for controlling flowering time | |
US5811536A (en) | Cauliflower floral meristem identity genes and methods of using same | |
WO1996034088A2 (en) | Control of floral induction in plants and uses therefor | |
CA2215335C (en) | Cauliflower floral meristem identity genes and methods of using same | |
AU697810B2 (en) | Genes regulating the response of (zea mays) to water deficit | |
AU3461099A (en) | Control of floral induction in plants and uses therefor | |
US7632984B2 (en) | Modulation of flowering time by the pft1 locus | |
Fan et al. | BnGF14-2c Positively Regulates Flowering via the Vernalization Pathway in Semi-Winter Rapeseed. Plants 2022, 11, 2312 | |
AU2012200697B2 (en) | Modification of plant and seed development and plant responses to stresses and stimuli (4) | |
AU2013202724A1 (en) | Modification of plant and seed development and plant responses to stresses and stimuli (6) |
Legal Events
Date | Code | Title | Description |
---|---|---|---|
AS | Assignment |
Owner name: SALK INSTITUTE FOR BIOLOGICAL STUDIES, THE, CALIFO Free format text: ASSIGNMENT OF ASSIGNORS INTEREST;ASSIGNORS:YANOFSKY, MARTIN;WEIGEL, DETLEF;REEL/FRAME:012058/0422;SIGNING DATES FROM 20010727 TO 20010730 |
|
AS | Assignment |
Owner name: REGENTS OF THE UNIVERSITY OF CALIFORNIA, THE, CALI Free format text: ASSIGNMENT OF ASSIGNORS INTEREST;ASSIGNOR:SALK INSTITUTE FOR BIOLOGICAL STUDIES, THE;REEL/FRAME:012200/0035 Effective date: 20010904 |
|
STCB | Information on status: application discontinuation |
Free format text: ABANDONED -- FAILURE TO RESPOND TO AN OFFICE ACTION |