CN112522259A - 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 - Google Patents
单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 Download PDFInfo
- Publication number
- CN112522259A CN112522259A CN202010996007.9A CN202010996007A CN112522259A CN 112522259 A CN112522259 A CN 112522259A CN 202010996007 A CN202010996007 A CN 202010996007A CN 112522259 A CN112522259 A CN 112522259A
- Authority
- CN
- China
- Prior art keywords
- gene
- editing
- seq
- vector
- oslg1
- 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.)
- Pending
Links
- 235000007164 Oryza sativa Nutrition 0.000 title claims abstract description 64
- 235000009566 rice Nutrition 0.000 title claims abstract description 64
- 241000196324 Embryophyta Species 0.000 title claims abstract description 54
- 238000000034 method Methods 0.000 title claims abstract description 40
- 239000000463 material Substances 0.000 title claims abstract description 16
- 240000007594 Oryza sativa Species 0.000 title 1
- 239000013598 vector Substances 0.000 claims abstract description 69
- 241000209094 Oryza Species 0.000 claims abstract description 68
- 238000010362 genome editing Methods 0.000 claims abstract description 62
- 108090000623 proteins and genes Proteins 0.000 claims abstract description 54
- 101710169836 Squamosa promoter-binding-like protein 8 Proteins 0.000 claims abstract description 43
- 108091028043 Nucleic acid sequence Proteins 0.000 claims abstract description 39
- 238000009395 breeding Methods 0.000 claims abstract description 21
- 230000001488 breeding effect Effects 0.000 claims abstract description 19
- 230000009261 transgenic effect Effects 0.000 claims abstract description 9
- 210000000349 chromosome Anatomy 0.000 claims abstract description 5
- 108091033319 polynucleotide Proteins 0.000 claims description 42
- 239000002157 polynucleotide Substances 0.000 claims description 42
- 102000040430 polynucleotide Human genes 0.000 claims description 42
- IAJOBQBIJHVGMQ-UHFFFAOYSA-N 2-amino-4-[hydroxy(methyl)phosphoryl]butanoic acid Chemical group CP(O)(=O)CCC(N)C(O)=O IAJOBQBIJHVGMQ-UHFFFAOYSA-N 0.000 claims description 13
- YQYJSBFKSSDGFO-UHFFFAOYSA-N Epihygromycin Natural products OC1C(O)C(C(=O)C)OC1OC(C(=C1)O)=CC=C1C=C(C)C(=O)NC1C(O)C(O)C2OCOC2C1O YQYJSBFKSSDGFO-UHFFFAOYSA-N 0.000 claims description 13
- 238000009396 hybridization Methods 0.000 claims description 12
- 239000003550 marker Substances 0.000 claims description 12
- 230000035772 mutation Effects 0.000 claims description 12
- 108091027544 Subgenomic mRNA Proteins 0.000 claims description 11
- 239000002773 nucleotide Substances 0.000 claims description 11
- 125000003729 nucleotide group Chemical group 0.000 claims description 11
- 238000012216 screening Methods 0.000 claims description 11
- 230000000694 effects Effects 0.000 claims description 8
- 230000001404 mediated effect Effects 0.000 claims description 8
- 238000006467 substitution reaction Methods 0.000 claims description 8
- 230000000295 complement effect Effects 0.000 claims description 7
- 230000002363 herbicidal effect Effects 0.000 claims description 7
- 238000012163 sequencing technique Methods 0.000 claims description 7
- 238000010367 cloning Methods 0.000 claims description 6
- 239000004009 herbicide Substances 0.000 claims description 6
- 238000003780 insertion Methods 0.000 claims description 6
- 230000037431 insertion Effects 0.000 claims description 6
- 230000008685 targeting Effects 0.000 claims description 6
- 102100035023 Carboxypeptidase B2 Human genes 0.000 claims description 5
- 108090000201 Carboxypeptidase B2 Proteins 0.000 claims description 5
- 238000012217 deletion Methods 0.000 claims description 5
- 230000037430 deletion Effects 0.000 claims description 5
- 108090000790 Enzymes Proteins 0.000 claims description 4
- 102000004190 Enzymes Human genes 0.000 claims description 4
- 238000005516 engineering process Methods 0.000 abstract description 12
- 230000006698 induction Effects 0.000 abstract description 9
- 230000008569 process Effects 0.000 abstract description 9
- 230000009466 transformation Effects 0.000 abstract description 9
- 230000002068 genetic effect Effects 0.000 abstract description 7
- 239000013077 target material Substances 0.000 abstract description 2
- 230000001939 inductive effect Effects 0.000 abstract 1
- 108091033409 CRISPR Proteins 0.000 description 21
- 210000004027 cell Anatomy 0.000 description 21
- 238000010354 CRISPR gene editing Methods 0.000 description 17
- 108020004414 DNA Proteins 0.000 description 11
- 239000000523 sample Substances 0.000 description 8
- 239000013604 expression vector Substances 0.000 description 7
- 102000004169 proteins and genes Human genes 0.000 description 7
- 125000000539 amino acid group Chemical group 0.000 description 6
- 150000007523 nucleic acids Chemical class 0.000 description 6
- 235000013339 cereals Nutrition 0.000 description 5
- 230000006872 improvement Effects 0.000 description 5
- 241000589158 Agrobacterium Species 0.000 description 4
- ZHNUHDYFZUAESO-UHFFFAOYSA-N Formamide Chemical compound NC=O ZHNUHDYFZUAESO-UHFFFAOYSA-N 0.000 description 4
- XKUKSGPZAADMRA-UHFFFAOYSA-N glycyl-glycyl-glycine Natural products NCC(=O)NCC(=O)NCC(O)=O XKUKSGPZAADMRA-UHFFFAOYSA-N 0.000 description 4
- 102000039446 nucleic acids Human genes 0.000 description 4
- 108020004707 nucleic acids Proteins 0.000 description 4
- 210000001519 tissue Anatomy 0.000 description 4
- 108091032973 (ribonucleotides)n+m Proteins 0.000 description 3
- 108091026890 Coding region Proteins 0.000 description 3
- 108020004705 Codon Proteins 0.000 description 3
- 239000005561 Glufosinate Substances 0.000 description 3
- 108010024078 alanyl-glycyl-serine Proteins 0.000 description 3
- 108010047857 aspartylglycine Proteins 0.000 description 3
- 230000001276 controlling effect Effects 0.000 description 3
- 230000006870 function Effects 0.000 description 3
- 108010050848 glycylleucine Proteins 0.000 description 3
- 238000004519 manufacturing process Methods 0.000 description 3
- 210000000056 organ Anatomy 0.000 description 3
- 108010031719 prolyl-serine Proteins 0.000 description 3
- 230000009467 reduction Effects 0.000 description 3
- 230000001105 regulatory effect Effects 0.000 description 3
- IAKHMKGGTNLKSZ-INIZCTEOSA-N (S)-colchicine Chemical compound C1([C@@H](NC(C)=O)CC2)=CC(=O)C(OC)=CC=C1C1=C2C=C(OC)C(OC)=C1OC IAKHMKGGTNLKSZ-INIZCTEOSA-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
- YYSWCHMLFJLLBJ-ZLUOBGJFSA-N Ala-Ala-Ser Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@@H](CO)C(O)=O YYSWCHMLFJLLBJ-ZLUOBGJFSA-N 0.000 description 2
- IJGRMHOSHXDMSA-UHFFFAOYSA-N Atomic nitrogen Chemical compound N#N IJGRMHOSHXDMSA-UHFFFAOYSA-N 0.000 description 2
- 206010064571 Gene mutation Diseases 0.000 description 2
- YJDALMUYJIENAG-QWRGUYRKSA-N Gly-Tyr-Asn Chemical compound C1=CC(=CC=C1C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)CN)O YJDALMUYJIENAG-QWRGUYRKSA-N 0.000 description 2
- 241000209510 Liliopsida Species 0.000 description 2
- YRAWWKUTNBILNT-FXQIFTODSA-N Met-Ala-Ala Chemical compound CSCC[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@@H](C)C(O)=O YRAWWKUTNBILNT-FXQIFTODSA-N 0.000 description 2
- 108091093037 Peptide nucleic acid Proteins 0.000 description 2
- RIYZXJVARWJLKS-KKUMJFAQSA-N Phe-Asp-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(O)=O)NC(=O)[C@@H](N)CC1=CC=CC=C1 RIYZXJVARWJLKS-KKUMJFAQSA-N 0.000 description 2
- LNICFEXCAHIJOR-DCAQKATOSA-N Pro-Ser-Leu Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O LNICFEXCAHIJOR-DCAQKATOSA-N 0.000 description 2
- FKNQFGJONOIPTF-UHFFFAOYSA-N Sodium cation Chemical compound [Na+] FKNQFGJONOIPTF-UHFFFAOYSA-N 0.000 description 2
- 108010076324 alanyl-glycyl-glycine Proteins 0.000 description 2
- 108010069020 alanyl-prolyl-glycine Proteins 0.000 description 2
- 230000009286 beneficial effect Effects 0.000 description 2
- 230000005540 biological transmission Effects 0.000 description 2
- 238000006243 chemical reaction Methods 0.000 description 2
- 108010016616 cysteinylglycine Proteins 0.000 description 2
- 238000013461 design Methods 0.000 description 2
- 210000005069 ears Anatomy 0.000 description 2
- 241001233957 eudicotyledons Species 0.000 description 2
- 108010078144 glutaminyl-glycine Proteins 0.000 description 2
- 239000000411 inducer Substances 0.000 description 2
- 238000005259 measurement Methods 0.000 description 2
- 239000013612 plasmid Substances 0.000 description 2
- 229920001184 polypeptide Polymers 0.000 description 2
- 108090000765 processed proteins & peptides Proteins 0.000 description 2
- 102000004196 processed proteins & peptides Human genes 0.000 description 2
- 238000011160 research Methods 0.000 description 2
- 150000003839 salts Chemical class 0.000 description 2
- 229910001415 sodium ion Inorganic materials 0.000 description 2
- 238000009423 ventilation Methods 0.000 description 2
- HKZAAJSTFUZYTO-LURJTMIESA-N (2s)-2-[[2-[[2-[[2-[(2-aminoacetyl)amino]acetyl]amino]acetyl]amino]acetyl]amino]-3-hydroxypropanoic acid Chemical compound NCC(=O)NCC(=O)NCC(=O)NCC(=O)N[C@@H](CO)C(O)=O HKZAAJSTFUZYTO-LURJTMIESA-N 0.000 description 1
- VGONTNSXDCQUGY-RRKCRQDMSA-N 2'-deoxyinosine Chemical group C1[C@H](O)[C@@H](CO)O[C@H]1N1C(N=CNC2=O)=C2N=C1 VGONTNSXDCQUGY-RRKCRQDMSA-N 0.000 description 1
- CXRCVCURMBFFOL-FXQIFTODSA-N Ala-Ala-Pro Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)N1CCC[C@H]1C(O)=O CXRCVCURMBFFOL-FXQIFTODSA-N 0.000 description 1
- JMCUQXTXLJEQSY-XKNYDFJKSA-N Ala-Asn-Asn-Pro Chemical compound C[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N1CCC[C@H]1C(O)=O JMCUQXTXLJEQSY-XKNYDFJKSA-N 0.000 description 1
- XQGIRPGAVLFKBJ-CIUDSAMLSA-N Ala-Asn-Lys Chemical compound N[C@@H](C)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCCN)C(=O)O XQGIRPGAVLFKBJ-CIUDSAMLSA-N 0.000 description 1
- 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 1
- XAGIMRPOEJSYER-CIUDSAMLSA-N Ala-Cys-Lys Chemical compound C[C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CCCCN)C(=O)O)N XAGIMRPOEJSYER-CIUDSAMLSA-N 0.000 description 1
- OILNWMNBLIHXQK-ZLUOBGJFSA-N Ala-Cys-Ser Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CS)C(=O)N[C@@H](CO)C(O)=O OILNWMNBLIHXQK-ZLUOBGJFSA-N 0.000 description 1
- WKOBSJOZRJJVRZ-FXQIFTODSA-N Ala-Glu-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O WKOBSJOZRJJVRZ-FXQIFTODSA-N 0.000 description 1
- LJFNNUBZSZCZFN-WHFBIAKZSA-N Ala-Gly-Cys Chemical compound N[C@@H](C)C(=O)NCC(=O)N[C@@H](CS)C(=O)O LJFNNUBZSZCZFN-WHFBIAKZSA-N 0.000 description 1
- 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 1
- ANGAOPNEPIDLPO-XVYDVKMFSA-N Ala-His-Cys Chemical compound C[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CS)C(=O)O)N ANGAOPNEPIDLPO-XVYDVKMFSA-N 0.000 description 1
- 108010076441 Ala-His-His Proteins 0.000 description 1
- HQJKCXHQNUCKMY-GHCJXIJMSA-N Ala-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](C)N HQJKCXHQNUCKMY-GHCJXIJMSA-N 0.000 description 1
- MEFILNJXAVSUTO-JXUBOQSCSA-N Ala-Leu-Thr Chemical compound C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O MEFILNJXAVSUTO-JXUBOQSCSA-N 0.000 description 1
- 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 1
- NINQYGGNRIBFSC-CIUDSAMLSA-N Ala-Lys-Ser Chemical compound NCCCC[C@H](NC(=O)[C@@H](N)C)C(=O)N[C@@H](CO)C(O)=O NINQYGGNRIBFSC-CIUDSAMLSA-N 0.000 description 1
- XUCHENWTTBFODJ-FXQIFTODSA-N Ala-Met-Ala Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](C)C(O)=O XUCHENWTTBFODJ-FXQIFTODSA-N 0.000 description 1
- GFEDXKNBZMPEDM-KZVJFYERSA-N Ala-Met-Thr Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O GFEDXKNBZMPEDM-KZVJFYERSA-N 0.000 description 1
- HYIDEIQUCBKIPL-CQDKDKBSSA-N Ala-Phe-His Chemical compound C[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)N HYIDEIQUCBKIPL-CQDKDKBSSA-N 0.000 description 1
- NZGRHTKZFSVPAN-BIIVOSGPSA-N Ala-Ser-Pro Chemical compound C[C@@H](C(=O)N[C@@H](CO)C(=O)N1CCC[C@@H]1C(=O)O)N NZGRHTKZFSVPAN-BIIVOSGPSA-N 0.000 description 1
- HCBKAOZYACJUEF-XQXXSGGOSA-N Ala-Thr-Gln Chemical compound N[C@@H](C)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@@H](CCC(N)=O)C(=O)O HCBKAOZYACJUEF-XQXXSGGOSA-N 0.000 description 1
- WNHNMKOFKCHKKD-BFHQHQDPSA-N Ala-Thr-Gly Chemical compound [H]N[C@@H](C)C(=O)N[C@@H]([C@@H](C)O)C(=O)NCC(O)=O WNHNMKOFKCHKKD-BFHQHQDPSA-N 0.000 description 1
- QOIGKCBMXUCDQU-KDXUFGMBSA-N Ala-Thr-Pro Chemical compound C[C@H]([C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](C)N)O QOIGKCBMXUCDQU-KDXUFGMBSA-N 0.000 description 1
- SGYSTDWPNPKJPP-GUBZILKMSA-N Arg-Ala-Arg Chemical compound NC(=N)NCCC[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O SGYSTDWPNPKJPP-GUBZILKMSA-N 0.000 description 1
- 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 1
- VYSRNGOMGHOJCK-GUBZILKMSA-N Arg-Ala-Met Chemical compound C[C@@H](C(=O)N[C@@H](CCSC)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N VYSRNGOMGHOJCK-GUBZILKMSA-N 0.000 description 1
- 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 1
- HJVGMOYJDDXLMI-AVGNSLFASA-N Arg-Arg-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CCCNC(N)=N)NC(=O)[C@@H](N)CCCNC(N)=N HJVGMOYJDDXLMI-AVGNSLFASA-N 0.000 description 1
- DCGLNNVKIZXQOJ-FXQIFTODSA-N Arg-Asn-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](CCCN=C(N)N)N DCGLNNVKIZXQOJ-FXQIFTODSA-N 0.000 description 1
- YSUVMPICYVWRBX-VEVYYDQMSA-N Arg-Asp-Thr Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O YSUVMPICYVWRBX-VEVYYDQMSA-N 0.000 description 1
- VNFWDYWTSHFRRG-SRVKXCTJSA-N Arg-Gln-Leu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O VNFWDYWTSHFRRG-SRVKXCTJSA-N 0.000 description 1
- CYXCAHZVPFREJD-LURJTMIESA-N Arg-Gly-Gly Chemical compound NC(=N)NCCC[C@H](N)C(=O)NCC(=O)NCC(O)=O CYXCAHZVPFREJD-LURJTMIESA-N 0.000 description 1
- HAVKMRGWNXMCDR-STQMWFEESA-N Arg-Gly-Phe Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)NCC(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O HAVKMRGWNXMCDR-STQMWFEESA-N 0.000 description 1
- YQGZIRIYGHNSQO-ZPFDUUQYSA-N Arg-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](CCCN=C(N)N)N YQGZIRIYGHNSQO-ZPFDUUQYSA-N 0.000 description 1
- FSNVAJOPUDVQAR-AVGNSLFASA-N Arg-Lys-Arg Chemical compound NC(=N)NCCC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O FSNVAJOPUDVQAR-AVGNSLFASA-N 0.000 description 1
- MTYLORHAQXVQOW-AVGNSLFASA-N Arg-Lys-Met Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCSC)C(O)=O MTYLORHAQXVQOW-AVGNSLFASA-N 0.000 description 1
- BSGSDLYGGHGMND-IHRRRGAJSA-N Arg-Phe-Cys Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N BSGSDLYGGHGMND-IHRRRGAJSA-N 0.000 description 1
- IGFJVXOATGZTHD-UHFFFAOYSA-N Arg-Phe-His Natural products NC(CCNC(=N)N)C(=O)NC(Cc1ccccc1)C(=O)NC(Cc2c[nH]cn2)C(=O)O IGFJVXOATGZTHD-UHFFFAOYSA-N 0.000 description 1
- UIUXXFIKWQVMEX-UFYCRDLUSA-N Arg-Phe-Tyr Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O UIUXXFIKWQVMEX-UFYCRDLUSA-N 0.000 description 1
- 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 1
- OGZBJJLRKQZRHL-KJEVXHAQSA-N Arg-Thr-Tyr Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 OGZBJJLRKQZRHL-KJEVXHAQSA-N 0.000 description 1
- XRLOBFSLPCHYLQ-ULQDDVLXSA-N Arg-Tyr-His Chemical compound C1=CC(=CC=C1C[C@@H](C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N)O XRLOBFSLPCHYLQ-ULQDDVLXSA-N 0.000 description 1
- HZPSDHRYYIORKR-WHFBIAKZSA-N Asn-Ala-Gly Chemical compound OC(=O)CNC(=O)[C@H](C)NC(=O)[C@@H](N)CC(N)=O HZPSDHRYYIORKR-WHFBIAKZSA-N 0.000 description 1
- GMRGSBAMMMVDGG-GUBZILKMSA-N Asn-Arg-Arg Chemical compound C(C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)NC(=O)[C@H](CC(=O)N)N)CN=C(N)N GMRGSBAMMMVDGG-GUBZILKMSA-N 0.000 description 1
- NKLRWRRVYGQNIH-GHCJXIJMSA-N Asn-Ile-Ala Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C)C(O)=O NKLRWRRVYGQNIH-GHCJXIJMSA-N 0.000 description 1
- GQRDIVQPSMPQME-ZPFDUUQYSA-N Asn-Ile-Leu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(C)C)C(O)=O GQRDIVQPSMPQME-ZPFDUUQYSA-N 0.000 description 1
- QDXQWFBLUVTOFL-FXQIFTODSA-N Asn-Met-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCSC)NC(=O)[C@H](CC(=O)N)N QDXQWFBLUVTOFL-FXQIFTODSA-N 0.000 description 1
- 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 1
- 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 1
- VPPXTHJNTYDNFJ-CIUDSAMLSA-N Asp-Ala-Lys Chemical compound C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CC(=O)O)N VPPXTHJNTYDNFJ-CIUDSAMLSA-N 0.000 description 1
- XPGVTUBABLRGHY-BIIVOSGPSA-N Asp-Ala-Pro Chemical compound C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CC(=O)O)N XPGVTUBABLRGHY-BIIVOSGPSA-N 0.000 description 1
- GWTLRDMPMJCNMH-WHFBIAKZSA-N Asp-Asn-Gly Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(N)=O)C(=O)NCC(O)=O GWTLRDMPMJCNMH-WHFBIAKZSA-N 0.000 description 1
- XACXDSRQIXRMNS-OLHMAJIHSA-N Asp-Asn-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](CC(=O)O)N)O XACXDSRQIXRMNS-OLHMAJIHSA-N 0.000 description 1
- NYQHSUGFEWDWPD-ACZMJKKPSA-N Asp-Gln-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC(=O)O)N NYQHSUGFEWDWPD-ACZMJKKPSA-N 0.000 description 1
- BKXPJCBEHWFSTF-ACZMJKKPSA-N Asp-Gln-Asp Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O BKXPJCBEHWFSTF-ACZMJKKPSA-N 0.000 description 1
- OVPHVTCDVYYTHN-AVGNSLFASA-N Asp-Glu-Phe Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 OVPHVTCDVYYTHN-AVGNSLFASA-N 0.000 description 1
- WYOSXGYAKZQPGF-SRVKXCTJSA-N Asp-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](CC(=O)O)N WYOSXGYAKZQPGF-SRVKXCTJSA-N 0.000 description 1
- CYCKJEFVFNRWEZ-UGYAYLCHSA-N Asp-Ile-Asn Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(N)=O)C(O)=O CYCKJEFVFNRWEZ-UGYAYLCHSA-N 0.000 description 1
- PYXXJFRXIYAESU-PCBIJLKTSA-N Asp-Ile-Phe Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O PYXXJFRXIYAESU-PCBIJLKTSA-N 0.000 description 1
- UMHUHHJMEXNSIV-CIUDSAMLSA-N Asp-Leu-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H](N)CC(O)=O UMHUHHJMEXNSIV-CIUDSAMLSA-N 0.000 description 1
- VSMYBNPOHYAXSD-GUBZILKMSA-N Asp-Lys-Glu Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O VSMYBNPOHYAXSD-GUBZILKMSA-N 0.000 description 1
- WOPJVEMFXYHZEE-SRVKXCTJSA-N Asp-Phe-Asp Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(O)=O)C(O)=O WOPJVEMFXYHZEE-SRVKXCTJSA-N 0.000 description 1
- UAXIKORUDGGIGA-DCAQKATOSA-N Asp-Pro-Lys Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CC(=O)O)N)C(=O)N[C@@H](CCCCN)C(=O)O UAXIKORUDGGIGA-DCAQKATOSA-N 0.000 description 1
- GCACQYDBDHRVGE-LKXGYXEUSA-N Asp-Thr-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H]([C@H](O)C)NC(=O)[C@@H](N)CC(O)=O GCACQYDBDHRVGE-LKXGYXEUSA-N 0.000 description 1
- 241000894006 Bacteria Species 0.000 description 1
- 201000004569 Blindness Diseases 0.000 description 1
- UKVGHFORADMBEN-GUBZILKMSA-N Cys-Arg-Arg Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O UKVGHFORADMBEN-GUBZILKMSA-N 0.000 description 1
- 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 1
- KCPOQGRVVXYLAC-KKUMJFAQSA-N Cys-Leu-Phe Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H](CS)N KCPOQGRVVXYLAC-KKUMJFAQSA-N 0.000 description 1
- BCFXQBXXDSEHRS-FXQIFTODSA-N Cys-Ser-Arg Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O BCFXQBXXDSEHRS-FXQIFTODSA-N 0.000 description 1
- RZSLYUUFFVHFRQ-FXQIFTODSA-N Gln-Ala-Glu Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(O)=O RZSLYUUFFVHFRQ-FXQIFTODSA-N 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
- LJEPDHWNQXPXMM-NHCYSSNCSA-N Gln-Arg-Val Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](C(C)C)C(O)=O LJEPDHWNQXPXMM-NHCYSSNCSA-N 0.000 description 1
- LPYPANUXJGFMGV-FXQIFTODSA-N Gln-Gln-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CCC(=O)N)N LPYPANUXJGFMGV-FXQIFTODSA-N 0.000 description 1
- RRBLZNIIMHSHQF-FXQIFTODSA-N Gln-Gln-Cys Chemical compound C(CC(=O)N)[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CS)C(=O)O)N RRBLZNIIMHSHQF-FXQIFTODSA-N 0.000 description 1
- 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 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
- KVXVVDFOZNYYKZ-DCAQKATOSA-N Gln-Gln-Leu Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O KVXVVDFOZNYYKZ-DCAQKATOSA-N 0.000 description 1
- NROSLUJMIQGFKS-IUCAKERBSA-N Gln-His-Gly Chemical compound C1=C(NC=N1)C[C@@H](C(=O)NCC(=O)O)NC(=O)[C@H](CCC(=O)N)N NROSLUJMIQGFKS-IUCAKERBSA-N 0.000 description 1
- ZNTDJIMJKNNSLR-RWRJDSDZSA-N Gln-Ile-Thr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)O)NC(=O)[C@H](CCC(=O)N)N ZNTDJIMJKNNSLR-RWRJDSDZSA-N 0.000 description 1
- CAXXTYYGFYTBPV-IUCAKERBSA-N Gln-Leu-Gly Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O CAXXTYYGFYTBPV-IUCAKERBSA-N 0.000 description 1
- LGWNISYVKDNJRP-FXQIFTODSA-N Gln-Ser-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCC(N)=O)C(O)=O LGWNISYVKDNJRP-FXQIFTODSA-N 0.000 description 1
- KVQOVQVGVKDZNW-GUBZILKMSA-N Gln-Ser-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CCC(=O)N)N KVQOVQVGVKDZNW-GUBZILKMSA-N 0.000 description 1
- LPIKVBWNNVFHCQ-GUBZILKMSA-N Gln-Ser-Leu Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O LPIKVBWNNVFHCQ-GUBZILKMSA-N 0.000 description 1
- GHAXJVNBAKGWEJ-AVGNSLFASA-N Gln-Ser-Tyr Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O GHAXJVNBAKGWEJ-AVGNSLFASA-N 0.000 description 1
- WVTIBGWZUMJBFY-GUBZILKMSA-N Glu-His-Ser Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CO)C(O)=O WVTIBGWZUMJBFY-GUBZILKMSA-N 0.000 description 1
- QJVZSVUYZFYLFQ-CIUDSAMLSA-N Glu-Pro-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](C)C(O)=O QJVZSVUYZFYLFQ-CIUDSAMLSA-N 0.000 description 1
- LSYFGBRDBIQYAQ-FHWLQOOXSA-N Glu-Tyr-Tyr Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O LSYFGBRDBIQYAQ-FHWLQOOXSA-N 0.000 description 1
- RMWAOBGCZZSJHE-UMNHJUIQSA-N Glu-Val-Pro Chemical compound CC(C)[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CCC(=O)O)N RMWAOBGCZZSJHE-UMNHJUIQSA-N 0.000 description 1
- SOYWRINXUSUWEQ-DLOVCJGASA-N Glu-Val-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CCC(O)=O SOYWRINXUSUWEQ-DLOVCJGASA-N 0.000 description 1
- UGVQELHRNUDMAA-BYPYZUCNSA-N Gly-Ala-Gly Chemical compound [NH3+]CC(=O)N[C@@H](C)C(=O)NCC([O-])=O UGVQELHRNUDMAA-BYPYZUCNSA-N 0.000 description 1
- JRDYDYXZKFNNRQ-XPUUQOCRSA-N Gly-Ala-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)CN JRDYDYXZKFNNRQ-XPUUQOCRSA-N 0.000 description 1
- KKBWDNZXYLGJEY-UHFFFAOYSA-N Gly-Arg-Pro Natural products NCC(=O)NC(CCNC(=N)N)C(=O)N1CCCC1C(=O)O KKBWDNZXYLGJEY-UHFFFAOYSA-N 0.000 description 1
- QGZSAHIZRQHCEQ-QWRGUYRKSA-N Gly-Asp-Tyr Chemical compound NCC(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 QGZSAHIZRQHCEQ-QWRGUYRKSA-N 0.000 description 1
- LGQZOQRDEUIZJY-YUMQZZPRSA-N Gly-Cys-Lys Chemical compound NCCCC[C@H](NC(=O)[C@H](CS)NC(=O)CN)C(O)=O LGQZOQRDEUIZJY-YUMQZZPRSA-N 0.000 description 1
- 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 1
- BEQGFMIBZFNROK-JGVFFNPUSA-N Gly-Glu-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCC(=O)O)NC(=O)CN)C(=O)O BEQGFMIBZFNROK-JGVFFNPUSA-N 0.000 description 1
- CCQOOWAONKGYKQ-BYPYZUCNSA-N Gly-Gly-Ala Chemical compound OC(=O)[C@H](C)NC(=O)CNC(=O)CN CCQOOWAONKGYKQ-BYPYZUCNSA-N 0.000 description 1
- KMSGYZQRXPUKGI-BYPYZUCNSA-N Gly-Gly-Asn Chemical compound NCC(=O)NCC(=O)N[C@H](C(O)=O)CC(N)=O KMSGYZQRXPUKGI-BYPYZUCNSA-N 0.000 description 1
- GDOZQTNZPCUARW-YFKPBYRVSA-N Gly-Gly-Glu Chemical compound NCC(=O)NCC(=O)N[C@H](C(O)=O)CCC(O)=O GDOZQTNZPCUARW-YFKPBYRVSA-N 0.000 description 1
- XPJBQTCXPJNIFE-ZETCQYMHSA-N Gly-Gly-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)CNC(=O)CN XPJBQTCXPJNIFE-ZETCQYMHSA-N 0.000 description 1
- IUZGUFAJDBHQQV-YUMQZZPRSA-N Gly-Leu-Asn Chemical compound NCC(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(N)=O)C(O)=O IUZGUFAJDBHQQV-YUMQZZPRSA-N 0.000 description 1
- VBOBNHSVQKKTOT-YUMQZZPRSA-N Gly-Lys-Ala Chemical compound [H]NCC(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O VBOBNHSVQKKTOT-YUMQZZPRSA-N 0.000 description 1
- PTIIBFKSLCYQBO-NHCYSSNCSA-N Gly-Lys-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)CN PTIIBFKSLCYQBO-NHCYSSNCSA-N 0.000 description 1
- MHZXESQPPXOING-KBPBESRZSA-N Gly-Lys-Phe Chemical compound [H]NCC(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O MHZXESQPPXOING-KBPBESRZSA-N 0.000 description 1
- DBJYVKDPGIFXFO-BQBZGAKWSA-N Gly-Met-Ala Chemical compound [H]NCC(=O)N[C@@H](CCSC)C(=O)N[C@@H](C)C(O)=O DBJYVKDPGIFXFO-BQBZGAKWSA-N 0.000 description 1
- MDKCBHZLQJZOCJ-STQMWFEESA-N Gly-Met-Tyr Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)O)NC(=O)CN MDKCBHZLQJZOCJ-STQMWFEESA-N 0.000 description 1
- WZSHYFGOLPXPLL-RYUDHWBXSA-N Gly-Phe-Glu Chemical compound NCC(=O)N[C@@H](Cc1ccccc1)C(=O)N[C@@H](CCC(O)=O)C(O)=O WZSHYFGOLPXPLL-RYUDHWBXSA-N 0.000 description 1
- LCRDMSSAKLTKBU-ZDLURKLDSA-N Gly-Ser-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)CN LCRDMSSAKLTKBU-ZDLURKLDSA-N 0.000 description 1
- GWCJMBNBFYBQCV-XPUUQOCRSA-N Gly-Val-Ala Chemical compound NCC(=O)N[C@@H](C(C)C)C(=O)N[C@@H](C)C(O)=O GWCJMBNBFYBQCV-XPUUQOCRSA-N 0.000 description 1
- JBCLFWXMTIKCCB-UHFFFAOYSA-N H-Gly-Phe-OH Natural products NCC(=O)NC(C(O)=O)CC1=CC=CC=C1 JBCLFWXMTIKCCB-UHFFFAOYSA-N 0.000 description 1
- 241000238631 Hexapoda Species 0.000 description 1
- BIAKMWKJMQLZOJ-ZKWXMUAHSA-N His-Ala-Ala Chemical compound C[C@H](NC(=O)[C@H](C)NC(=O)[C@@H](N)Cc1cnc[nH]1)C(O)=O BIAKMWKJMQLZOJ-ZKWXMUAHSA-N 0.000 description 1
- WMKXFMUJRCEGRP-SRVKXCTJSA-N His-Asn-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)N WMKXFMUJRCEGRP-SRVKXCTJSA-N 0.000 description 1
- UOAVQQRILDGZEN-SRVKXCTJSA-N His-Asp-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O UOAVQQRILDGZEN-SRVKXCTJSA-N 0.000 description 1
- LCNNHVQNFNJLGK-AVGNSLFASA-N His-Gln-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)N LCNNHVQNFNJLGK-AVGNSLFASA-N 0.000 description 1
- PYNUBZSXKQKAHL-UWVGGRQHSA-N His-Gly-Arg Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)NCC(=O)N[C@@H](CCCNC(N)=N)C(O)=O PYNUBZSXKQKAHL-UWVGGRQHSA-N 0.000 description 1
- JIUYRPFQJJRSJB-QWRGUYRKSA-N His-His-Gly Chemical compound C([C@H](N)C(=O)N[C@@H](CC=1NC=NC=1)C(=O)NCC(O)=O)C1=CN=CN1 JIUYRPFQJJRSJB-QWRGUYRKSA-N 0.000 description 1
- JBSLJUPMTYLLFH-MELADBBJSA-N His-His-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC2=CN=CN2)NC(=O)[C@H](CC3=CN=CN3)N)C(=O)O JBSLJUPMTYLLFH-MELADBBJSA-N 0.000 description 1
- BZKDJRSZWLPJNI-SRVKXCTJSA-N His-His-Ser Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CO)C(O)=O BZKDJRSZWLPJNI-SRVKXCTJSA-N 0.000 description 1
- YAALVYQFVJNXIV-KKUMJFAQSA-N His-Leu-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H](N)CC1=CN=CN1 YAALVYQFVJNXIV-KKUMJFAQSA-N 0.000 description 1
- FJCGVRRVBKYYOU-DCAQKATOSA-N His-Met-Ser Chemical compound CSCC[C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](CC1=CN=CN1)N FJCGVRRVBKYYOU-DCAQKATOSA-N 0.000 description 1
- STGQSBKUYSPPIG-CIUDSAMLSA-N His-Ser-Asp Chemical compound OC(=O)C[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@@H](N)CC1=CN=CN1 STGQSBKUYSPPIG-CIUDSAMLSA-N 0.000 description 1
- DAKSMIWQZPHRIB-BZSNNMDCSA-N His-Tyr-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(C)C)C(O)=O DAKSMIWQZPHRIB-BZSNNMDCSA-N 0.000 description 1
- 206010020649 Hyperkeratosis Diseases 0.000 description 1
- YPWHUFAAMNHMGS-QSFUFRPTSA-N Ile-Ala-His Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](C)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N YPWHUFAAMNHMGS-QSFUFRPTSA-N 0.000 description 1
- QLRMMMQNCWBNPQ-QXEWZRGKSA-N Ile-Arg-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)NCC(=O)O)N QLRMMMQNCWBNPQ-QXEWZRGKSA-N 0.000 description 1
- NKRJALPCDNXULF-BYULHYEWSA-N Ile-Asp-Gly Chemical compound [H]N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(O)=O)C(=O)NCC(O)=O NKRJALPCDNXULF-BYULHYEWSA-N 0.000 description 1
- WTOAPTKSZJJWKK-HTFCKZLJSA-N Ile-Cys-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H]([C@@H](C)CC)C(=O)O)N WTOAPTKSZJJWKK-HTFCKZLJSA-N 0.000 description 1
- BSWLQVGEVFYGIM-ZPFDUUQYSA-N Ile-Gln-Arg Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)N BSWLQVGEVFYGIM-ZPFDUUQYSA-N 0.000 description 1
- KFVUBLZRFSVDGO-BYULHYEWSA-N Ile-Gly-Asp Chemical compound CC[C@H](C)[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CC(O)=O KFVUBLZRFSVDGO-BYULHYEWSA-N 0.000 description 1
- HYLIOBDWPQNLKI-HVTMNAMFSA-N Ile-His-Gln Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N HYLIOBDWPQNLKI-HVTMNAMFSA-N 0.000 description 1
- FGBRXCZYVRFNKQ-MXAVVETBSA-N Ile-Phe-Ser Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CO)C(=O)O)N FGBRXCZYVRFNKQ-MXAVVETBSA-N 0.000 description 1
- CNMOKANDJMLAIF-CIQUZCHMSA-N Ile-Thr-Ala Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C)C(O)=O CNMOKANDJMLAIF-CIQUZCHMSA-N 0.000 description 1
- 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 1
- HGCNKOLVKRAVHD-UHFFFAOYSA-N L-Met-L-Phe Natural products CSCCC(N)C(=O)NC(C(O)=O)CC1=CC=CC=C1 HGCNKOLVKRAVHD-UHFFFAOYSA-N 0.000 description 1
- UGTHTQWIQKEDEH-BQBZGAKWSA-N L-alanyl-L-prolylglycine zwitterion Chemical compound C[C@H](N)C(=O)N1CCC[C@H]1C(=O)NCC(O)=O UGTHTQWIQKEDEH-BQBZGAKWSA-N 0.000 description 1
- SENJXOPIZNYLHU-UHFFFAOYSA-N L-leucyl-L-arginine Natural products CC(C)CC(N)C(=O)NC(C(O)=O)CCCN=C(N)N SENJXOPIZNYLHU-UHFFFAOYSA-N 0.000 description 1
- HBJZFCIVFIBNSV-DCAQKATOSA-N Leu-Arg-Asn Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC(N)=O)C(O)=O HBJZFCIVFIBNSV-DCAQKATOSA-N 0.000 description 1
- GLBNEGIOFRVRHO-JYJNAYRXSA-N Leu-Gln-Phe Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O GLBNEGIOFRVRHO-JYJNAYRXSA-N 0.000 description 1
- OGUUKPXUTHOIAV-SDDRHHMPSA-N Leu-Glu-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N OGUUKPXUTHOIAV-SDDRHHMPSA-N 0.000 description 1
- BABSVXFGKFLIGW-UWVGGRQHSA-N Leu-Gly-Arg Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCNC(N)=N BABSVXFGKFLIGW-UWVGGRQHSA-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
- UCDHVOALNXENLC-KBPBESRZSA-N Leu-Gly-Tyr Chemical compound CC(C)C[C@H]([NH3+])C(=O)NCC(=O)N[C@H](C([O-])=O)CC1=CC=C(O)C=C1 UCDHVOALNXENLC-KBPBESRZSA-N 0.000 description 1
- BKTXKJMNTSMJDQ-AVGNSLFASA-N Leu-His-Gln Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N BKTXKJMNTSMJDQ-AVGNSLFASA-N 0.000 description 1
- CFZZDVMBRYFFNU-QWRGUYRKSA-N Leu-His-Gly Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CNC=N1)C(=O)NCC(O)=O CFZZDVMBRYFFNU-QWRGUYRKSA-N 0.000 description 1
- XVZCXCTYGHPNEM-UHFFFAOYSA-N Leu-Leu-Pro Natural products CC(C)CC(N)C(=O)NC(CC(C)C)C(=O)N1CCCC1C(O)=O XVZCXCTYGHPNEM-UHFFFAOYSA-N 0.000 description 1
- YUTNOGOMBNYPFH-XUXIUFHCSA-N Leu-Pro-Ile Chemical compound [H]N[C@@H](CC(C)C)C(=O)N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)CC)C(O)=O YUTNOGOMBNYPFH-XUXIUFHCSA-N 0.000 description 1
- IZPVWNSAVUQBGP-CIUDSAMLSA-N Leu-Ser-Asp Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(O)=O)C(O)=O IZPVWNSAVUQBGP-CIUDSAMLSA-N 0.000 description 1
- AXVIGSRGTMNSJU-YESZJQIVSA-N Leu-Tyr-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N2CCC[C@@H]2C(=O)O)N AXVIGSRGTMNSJU-YESZJQIVSA-N 0.000 description 1
- CGHXMODRYJISSK-NHCYSSNCSA-N Leu-Val-Asp Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CC(O)=O CGHXMODRYJISSK-NHCYSSNCSA-N 0.000 description 1
- XFIHDSBIPWEYJJ-YUMQZZPRSA-N Lys-Ala-Gly Chemical compound OC(=O)CNC(=O)[C@H](C)NC(=O)[C@@H](N)CCCCN XFIHDSBIPWEYJJ-YUMQZZPRSA-N 0.000 description 1
- 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 1
- ZTPWXNOOKAXPPE-DCAQKATOSA-N Lys-Arg-Cys Chemical compound C(CCN)C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CS)C(=O)O)N ZTPWXNOOKAXPPE-DCAQKATOSA-N 0.000 description 1
- DGAAQRAUOFHBFJ-CIUDSAMLSA-N Lys-Asn-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](C)C(O)=O DGAAQRAUOFHBFJ-CIUDSAMLSA-N 0.000 description 1
- HIIZIQUUHIXUJY-GUBZILKMSA-N Lys-Asp-Gln Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O HIIZIQUUHIXUJY-GUBZILKMSA-N 0.000 description 1
- 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 1
- JZMGVXLDOQOKAH-UWVGGRQHSA-N Lys-Gly-Met Chemical compound [H]N[C@@H](CCCCN)C(=O)NCC(=O)N[C@@H](CCSC)C(O)=O JZMGVXLDOQOKAH-UWVGGRQHSA-N 0.000 description 1
- 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 1
- CTJUSALVKAWFFU-CIUDSAMLSA-N Lys-Ser-Cys Chemical compound C(CCN)C[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CS)C(=O)O)N CTJUSALVKAWFFU-CIUDSAMLSA-N 0.000 description 1
- MIMXMVDLMDMOJD-BZSNNMDCSA-N Lys-Tyr-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(C)C)C(O)=O MIMXMVDLMDMOJD-BZSNNMDCSA-N 0.000 description 1
- OHXUUQDOBQKSNB-AVGNSLFASA-N Lys-Val-Arg Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O OHXUUQDOBQKSNB-AVGNSLFASA-N 0.000 description 1
- QFSYGUMEANRNJE-DCAQKATOSA-N Lys-Val-Cys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CCCCN)N QFSYGUMEANRNJE-DCAQKATOSA-N 0.000 description 1
- OSOLWRWQADPDIQ-DCAQKATOSA-N Met-Asp-Leu Chemical compound [H]N[C@@H](CCSC)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O OSOLWRWQADPDIQ-DCAQKATOSA-N 0.000 description 1
- 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 1
- 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 1
- YLDSJJOGQNEQJK-AVGNSLFASA-N Met-Pro-Leu Chemical compound CSCC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC(C)C)C(O)=O YLDSJJOGQNEQJK-AVGNSLFASA-N 0.000 description 1
- LPNWWHBFXPNHJG-AVGNSLFASA-N Met-Val-Lys Chemical compound CSCC[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CCCCN LPNWWHBFXPNHJG-AVGNSLFASA-N 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
- KZNQNBZMBZJQJO-UHFFFAOYSA-N N-glycyl-L-proline Natural products NCC(=O)N1CCCC1C(O)=O KZNQNBZMBZJQJO-UHFFFAOYSA-N 0.000 description 1
- 108010002311 N-glycylglutamic acid Proteins 0.000 description 1
- 108091034117 Oligonucleotide Proteins 0.000 description 1
- 208000012868 Overgrowth Diseases 0.000 description 1
- NKLDZIPTGKBDBB-HTUGSXCWSA-N Phe-Gln-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC1=CC=CC=C1)N)O NKLDZIPTGKBDBB-HTUGSXCWSA-N 0.000 description 1
- CZQZSMJXFGGBHM-KKUMJFAQSA-N Phe-Pro-Gln Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CC2=CC=CC=C2)N)C(=O)N[C@@H](CCC(=O)N)C(=O)O CZQZSMJXFGGBHM-KKUMJFAQSA-N 0.000 description 1
- CKJACGQPCPMWIT-UFYCRDLUSA-N Phe-Pro-Phe Chemical compound C([C@H](N)C(=O)N1[C@@H](CCC1)C(=O)N[C@@H](CC=1C=CC=CC=1)C(O)=O)C1=CC=CC=C1 CKJACGQPCPMWIT-UFYCRDLUSA-N 0.000 description 1
- ZJPGOXWRFNKIQL-JYJNAYRXSA-N Phe-Pro-Pro Chemical compound C([C@H](N)C(=O)N1[C@@H](CCC1)C(=O)N1[C@@H](CCC1)C(O)=O)C1=CC=CC=C1 ZJPGOXWRFNKIQL-JYJNAYRXSA-N 0.000 description 1
- XOHJOMKCRLHGCY-UNQGMJICSA-N Phe-Pro-Thr Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)O)C(O)=O XOHJOMKCRLHGCY-UNQGMJICSA-N 0.000 description 1
- OLHDPZMYUSBGDE-GUBZILKMSA-N Pro-Arg-Cys Chemical compound C1C[C@H](NC1)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CS)C(=O)O OLHDPZMYUSBGDE-GUBZILKMSA-N 0.000 description 1
- LNOWDSPAYBWJOR-PEDHHIEDSA-N Pro-Ile-Ile Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O LNOWDSPAYBWJOR-PEDHHIEDSA-N 0.000 description 1
- BRJGUPWVFXKBQI-XUXIUFHCSA-N Pro-Leu-Ile Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O BRJGUPWVFXKBQI-XUXIUFHCSA-N 0.000 description 1
- DRKAXLDECUGLFE-ULQDDVLXSA-N Pro-Leu-Phe Chemical compound CC(C)C[C@H](NC(=O)[C@@H]1CCCN1)C(=O)N[C@@H](Cc1ccccc1)C(O)=O DRKAXLDECUGLFE-ULQDDVLXSA-N 0.000 description 1
- 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 1
- 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 1
- MKGIILKDUGDRRO-FXQIFTODSA-N Pro-Ser-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@@H]1CCCN1 MKGIILKDUGDRRO-FXQIFTODSA-N 0.000 description 1
- 244000184734 Pyrus japonica Species 0.000 description 1
- 108091028664 Ribonucleotide Proteins 0.000 description 1
- ZUGXSSFMTXKHJS-ZLUOBGJFSA-N Ser-Ala-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](C)C(=O)N[C@@H](C)C(O)=O ZUGXSSFMTXKHJS-ZLUOBGJFSA-N 0.000 description 1
- WTUJZHKANPDPIN-CIUDSAMLSA-N Ser-Ala-Lys Chemical compound C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CO)N WTUJZHKANPDPIN-CIUDSAMLSA-N 0.000 description 1
- BRKHVZNDAOMAHX-BIIVOSGPSA-N Ser-Ala-Pro Chemical compound C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CO)N BRKHVZNDAOMAHX-BIIVOSGPSA-N 0.000 description 1
- QGMLKFGTGXWAHF-IHRRRGAJSA-N Ser-Arg-Phe Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O QGMLKFGTGXWAHF-IHRRRGAJSA-N 0.000 description 1
- 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 1
- CTRHXXXHUJTTRZ-ZLUOBGJFSA-N Ser-Asp-Cys Chemical compound C([C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CO)N)C(=O)O CTRHXXXHUJTTRZ-ZLUOBGJFSA-N 0.000 description 1
- VAIZFHMTBFYJIA-ACZMJKKPSA-N Ser-Asp-Gln Chemical compound OC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CCC(N)=O VAIZFHMTBFYJIA-ACZMJKKPSA-N 0.000 description 1
- BNFVPSRLHHPQKS-WHFBIAKZSA-N Ser-Asp-Gly Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(O)=O)C(=O)NCC(O)=O BNFVPSRLHHPQKS-WHFBIAKZSA-N 0.000 description 1
- KNCJWSPMTFFJII-ZLUOBGJFSA-N Ser-Cys-Asp Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CS)C(=O)N[C@@H](CC(O)=O)C(O)=O KNCJWSPMTFFJII-ZLUOBGJFSA-N 0.000 description 1
- UOLGINIHBRIECN-FXQIFTODSA-N Ser-Glu-Glu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O UOLGINIHBRIECN-FXQIFTODSA-N 0.000 description 1
- UIGMAMGZOJVTDN-WHFBIAKZSA-N Ser-Gly-Ser Chemical compound OC[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O UIGMAMGZOJVTDN-WHFBIAKZSA-N 0.000 description 1
- XXXAXOWMBOKTRN-XPUUQOCRSA-N Ser-Gly-Val Chemical compound [H]N[C@@H](CO)C(=O)NCC(=O)N[C@@H](C(C)C)C(O)=O XXXAXOWMBOKTRN-XPUUQOCRSA-N 0.000 description 1
- MOQDPPUMFSMYOM-KKUMJFAQSA-N Ser-His-Phe Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)O)NC(=O)[C@H](CC2=CN=CN2)NC(=O)[C@H](CO)N MOQDPPUMFSMYOM-KKUMJFAQSA-N 0.000 description 1
- PTWIYDNFWPXQSD-GARJFASQSA-N Ser-Lys-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCCCN)NC(=O)[C@H](CO)N)C(=O)O PTWIYDNFWPXQSD-GARJFASQSA-N 0.000 description 1
- ZGFRMNZZTOVBOU-CIUDSAMLSA-N Ser-Met-Gln Chemical compound N[C@@H](CO)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCC(N)=O)C(=O)O ZGFRMNZZTOVBOU-CIUDSAMLSA-N 0.000 description 1
- 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 1
- PPCZVWHJWJFTFN-ZLUOBGJFSA-N Ser-Ser-Asp Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(O)=O)C(O)=O PPCZVWHJWJFTFN-ZLUOBGJFSA-N 0.000 description 1
- BMKNXTJLHFIAAH-CIUDSAMLSA-N Ser-Ser-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O BMKNXTJLHFIAAH-CIUDSAMLSA-N 0.000 description 1
- OZPDGESCTGGNAD-CIUDSAMLSA-N Ser-Ser-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@@H](N)CO OZPDGESCTGGNAD-CIUDSAMLSA-N 0.000 description 1
- PYTKULIABVRXSC-BWBBJGPYSA-N Ser-Ser-Thr Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(O)=O PYTKULIABVRXSC-BWBBJGPYSA-N 0.000 description 1
- DYEGLQRVMBWQLD-IXOXFDKPSA-N Ser-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](CO)N)O DYEGLQRVMBWQLD-IXOXFDKPSA-N 0.000 description 1
- 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 1
- VVKVHAOOUGNDPJ-SRVKXCTJSA-N Ser-Tyr-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CO)C(O)=O VVKVHAOOUGNDPJ-SRVKXCTJSA-N 0.000 description 1
- RYYWUUFWQRZTIU-UHFFFAOYSA-N Thiophosphoric acid Chemical class OP(O)(S)=O RYYWUUFWQRZTIU-UHFFFAOYSA-N 0.000 description 1
- 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 1
- MQCPGOZXFSYJPS-KZVJFYERSA-N Thr-Ala-Arg Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O MQCPGOZXFSYJPS-KZVJFYERSA-N 0.000 description 1
- JNQZPAWOPBZGIX-RCWTZXSCSA-N Thr-Arg-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)[C@@H](C)O)CCCN=C(N)N JNQZPAWOPBZGIX-RCWTZXSCSA-N 0.000 description 1
- YOSLMIPKOUAHKI-OLHMAJIHSA-N Thr-Asp-Asp Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O YOSLMIPKOUAHKI-OLHMAJIHSA-N 0.000 description 1
- DHPPWTOLRWYIDS-XKBZYTNZSA-N Thr-Cys-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CS)C(=O)N[C@@H](CCC(O)=O)C(O)=O DHPPWTOLRWYIDS-XKBZYTNZSA-N 0.000 description 1
- RKDFEMGVMMYYNG-WDCWCFNPSA-N Thr-Gln-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O RKDFEMGVMMYYNG-WDCWCFNPSA-N 0.000 description 1
- MEJHFIOYJHTWMK-VOAKCMCISA-N Thr-Leu-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H](N)[C@@H](C)O MEJHFIOYJHTWMK-VOAKCMCISA-N 0.000 description 1
- ZSPQUTWLWGWTPS-HJGDQZAQSA-N Thr-Lys-Asp Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(O)=O)C(O)=O ZSPQUTWLWGWTPS-HJGDQZAQSA-N 0.000 description 1
- GUHLYMZJVXUIPO-RCWTZXSCSA-N Thr-Met-Val Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](C(C)C)C(O)=O GUHLYMZJVXUIPO-RCWTZXSCSA-N 0.000 description 1
- RVMNUBQWPVOUKH-HEIBUPTGSA-N Thr-Ser-Thr Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(O)=O RVMNUBQWPVOUKH-HEIBUPTGSA-N 0.000 description 1
- BZTSQFWJNJYZSX-JRQIVUDYSA-N Thr-Tyr-Asp Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(O)=O)C(O)=O BZTSQFWJNJYZSX-JRQIVUDYSA-N 0.000 description 1
- PELIQFPESHBTMA-WLTAIBSBSA-N Thr-Tyr-Gly Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@H](C(=O)NCC(O)=O)CC1=CC=C(O)C=C1 PELIQFPESHBTMA-WLTAIBSBSA-N 0.000 description 1
- JAWUQFCGNVEDRN-MEYUZBJRSA-N Thr-Tyr-Leu Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N[C@@H](CC(C)C)C(=O)O)N)O JAWUQFCGNVEDRN-MEYUZBJRSA-N 0.000 description 1
- FYBFTPLPAXZBOY-KKHAAJSZSA-N Thr-Val-Asp Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O FYBFTPLPAXZBOY-KKHAAJSZSA-N 0.000 description 1
- 108700019146 Transgenes Proteins 0.000 description 1
- DNUJCLUFRGGSDJ-YLVFBTJISA-N Trp-Gly-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CC1=CNC2=CC=CC=C21)N DNUJCLUFRGGSDJ-YLVFBTJISA-N 0.000 description 1
- GFHYISDTIWZUSU-QWRGUYRKSA-N Tyr-Asn-Gly Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(N)=O)C(=O)NCC(O)=O GFHYISDTIWZUSU-QWRGUYRKSA-N 0.000 description 1
- QUILOGWWLXMSAT-IHRRRGAJSA-N Tyr-Gln-Gln Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O QUILOGWWLXMSAT-IHRRRGAJSA-N 0.000 description 1
- XGZBEGGGAUQBMB-KJEVXHAQSA-N Tyr-Pro-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@@H]1CCCN1C(=O)[C@H](CC2=CC=C(C=C2)O)N)O XGZBEGGGAUQBMB-KJEVXHAQSA-N 0.000 description 1
- CWOSXNKDOACNJN-BZSNNMDCSA-N Val-Arg-Trp Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)O)N CWOSXNKDOACNJN-BZSNNMDCSA-N 0.000 description 1
- QHFQQRKNGCXTHL-AUTRQRHGSA-N Val-Gln-Glu Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O QHFQQRKNGCXTHL-AUTRQRHGSA-N 0.000 description 1
- KZKMBGXCNLPYKD-YEPSODPASA-N Val-Gly-Thr Chemical compound CC(C)[C@H](N)C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(O)=O KZKMBGXCNLPYKD-YEPSODPASA-N 0.000 description 1
- BTWMICVCQLKKNR-DCAQKATOSA-N Val-Leu-Ser Chemical compound CC(C)[C@H]([NH3+])C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C([O-])=O BTWMICVCQLKKNR-DCAQKATOSA-N 0.000 description 1
- CXWJFWAZIVWBOS-XQQFMLRXSA-N Val-Lys-Pro Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N1CCC[C@@H]1C(=O)O)N CXWJFWAZIVWBOS-XQQFMLRXSA-N 0.000 description 1
- RSGHLMMKXJGCMK-JYJNAYRXSA-N Val-Met-Phe Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)N RSGHLMMKXJGCMK-JYJNAYRXSA-N 0.000 description 1
- SSYBNWFXCFNRFN-GUBZILKMSA-N Val-Pro-Ser Chemical compound CC(C)[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O SSYBNWFXCFNRFN-GUBZILKMSA-N 0.000 description 1
- PZTZYZUTCPZWJH-FXQIFTODSA-N Val-Ser-Ser Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)O)N PZTZYZUTCPZWJH-FXQIFTODSA-N 0.000 description 1
- AEFJNECXZCODJM-UWVGGRQHSA-N Val-Val-Gly Chemical compound CC(C)[C@H]([NH3+])C(=O)N[C@@H](C(C)C)C(=O)NCC([O-])=O AEFJNECXZCODJM-UWVGGRQHSA-N 0.000 description 1
- JVGDAEKKZKKZFO-RCWTZXSCSA-N Val-Val-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](C(C)C)NC(=O)[C@H](C(C)C)N)O JVGDAEKKZKKZFO-RCWTZXSCSA-N 0.000 description 1
- 241000607479 Yersinia pestis Species 0.000 description 1
- 230000002411 adverse Effects 0.000 description 1
- 108010041407 alanylaspartic acid Proteins 0.000 description 1
- 108010044940 alanylglutamine Proteins 0.000 description 1
- 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 1
- 230000004075 alteration Effects 0.000 description 1
- 238000004458 analytical method Methods 0.000 description 1
- 230000000692 anti-sense effect Effects 0.000 description 1
- 108010013835 arginine glutamate Proteins 0.000 description 1
- 108010008355 arginyl-glutamine Proteins 0.000 description 1
- 108010043240 arginyl-leucyl-glycine Proteins 0.000 description 1
- 108010068380 arginylarginine Proteins 0.000 description 1
- 108010077245 asparaginyl-proline Proteins 0.000 description 1
- 108010040443 aspartyl-aspartic acid Proteins 0.000 description 1
- 230000008901 benefit Effects 0.000 description 1
- 102000005936 beta-Galactosidase Human genes 0.000 description 1
- 108010005774 beta-Galactosidase Proteins 0.000 description 1
- 230000003115 biocidal effect Effects 0.000 description 1
- 239000000969 carrier Substances 0.000 description 1
- 230000008859 change Effects 0.000 description 1
- 239000003795 chemical substances by application Substances 0.000 description 1
- 238000003776 cleavage reaction Methods 0.000 description 1
- 229960001338 colchicine Drugs 0.000 description 1
- 150000001875 compounds Chemical class 0.000 description 1
- 238000010276 construction Methods 0.000 description 1
- 238000007796 conventional method Methods 0.000 description 1
- 238000012272 crop production Methods 0.000 description 1
- 238000012364 cultivation method Methods 0.000 description 1
- 238000012258 culturing Methods 0.000 description 1
- 239000005549 deoxyribonucleoside Substances 0.000 description 1
- 239000005547 deoxyribonucleotide Substances 0.000 description 1
- 125000002637 deoxyribonucleotide group Chemical group 0.000 description 1
- 230000001419 dependent effect Effects 0.000 description 1
- 230000000368 destabilizing effect Effects 0.000 description 1
- 238000011161 development Methods 0.000 description 1
- 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 1
- 201000010099 disease Diseases 0.000 description 1
- 208000037265 diseases, disorders, signs and symptoms Diseases 0.000 description 1
- 238000004520 electroporation Methods 0.000 description 1
- 230000007613 environmental effect Effects 0.000 description 1
- 210000003527 eukaryotic cell Anatomy 0.000 description 1
- 239000003337 fertilizer Substances 0.000 description 1
- 238000000684 flow cytometry Methods 0.000 description 1
- 102000034287 fluorescent proteins Human genes 0.000 description 1
- 108091006047 fluorescent proteins Proteins 0.000 description 1
- 231100000221 frame shift mutation induction Toxicity 0.000 description 1
- 230000037433 frameshift Effects 0.000 description 1
- 108010055341 glutamyl-glutamic acid Proteins 0.000 description 1
- 108010049041 glutamylalanine Proteins 0.000 description 1
- 108010072405 glycyl-aspartyl-glycine Proteins 0.000 description 1
- 108010067216 glycyl-glycyl-glycine Proteins 0.000 description 1
- 108010010096 glycyl-glycyl-tyrosine Proteins 0.000 description 1
- 108010020688 glycylhistidine Proteins 0.000 description 1
- 108010015792 glycyllysine Proteins 0.000 description 1
- 108010077515 glycylproline Proteins 0.000 description 1
- 108010028295 histidylhistidine Proteins 0.000 description 1
- 108010018006 histidylserine Proteins 0.000 description 1
- 108010000761 leucylarginine Proteins 0.000 description 1
- 108010057821 leucylproline Proteins 0.000 description 1
- 108010064235 lysylglycine Proteins 0.000 description 1
- 108010016686 methionyl-alanyl-serine Proteins 0.000 description 1
- 108010068488 methionylphenylalanine Proteins 0.000 description 1
- 238000000520 microinjection Methods 0.000 description 1
- 239000000203 mixture Substances 0.000 description 1
- 238000012986 modification Methods 0.000 description 1
- 230000004048 modification Effects 0.000 description 1
- 238000002703 mutagenesis Methods 0.000 description 1
- 231100000350 mutagenesis Toxicity 0.000 description 1
- 229910052757 nitrogen Inorganic materials 0.000 description 1
- 239000000575 pesticide Substances 0.000 description 1
- 150000008298 phosphoramidates Chemical class 0.000 description 1
- 230000029553 photosynthesis Effects 0.000 description 1
- 238000010672 photosynthesis Methods 0.000 description 1
- 229920000642 polymer Polymers 0.000 description 1
- 210000001236 prokaryotic cell Anatomy 0.000 description 1
- 108010077112 prolyl-proline Proteins 0.000 description 1
- 108010029020 prolylglycine Proteins 0.000 description 1
- 239000002342 ribonucleoside Substances 0.000 description 1
- 239000002336 ribonucleotide Substances 0.000 description 1
- 125000002652 ribonucleotide group Chemical group 0.000 description 1
- 230000007017 scission Effects 0.000 description 1
- 108010069117 seryl-lysyl-aspartic acid Proteins 0.000 description 1
- 108010071207 serylmethionine Proteins 0.000 description 1
- 238000002741 site-directed mutagenesis Methods 0.000 description 1
- 238000009331 sowing Methods 0.000 description 1
- 241000894007 species Species 0.000 description 1
- 239000000126 substance Substances 0.000 description 1
- 238000003786 synthesis reaction Methods 0.000 description 1
- 238000012360 testing method Methods 0.000 description 1
- 108010061238 threonyl-glycine Proteins 0.000 description 1
- 238000013518 transcription Methods 0.000 description 1
- 230000035897 transcription Effects 0.000 description 1
- 238000010361 transduction Methods 0.000 description 1
- 230000026683 transduction Effects 0.000 description 1
- 238000012546 transfer Methods 0.000 description 1
- 238000011426 transformation method Methods 0.000 description 1
- 230000001131 transforming effect Effects 0.000 description 1
- 230000001052 transient effect Effects 0.000 description 1
- 238000013519 translation Methods 0.000 description 1
- 108010080629 tryptophan-leucine Proteins 0.000 description 1
- 108010035534 tyrosyl-leucyl-alanine Proteins 0.000 description 1
- 108010077037 tyrosyl-tyrosyl-phenylalanine Proteins 0.000 description 1
- IBIDRSSEHFLGSD-UHFFFAOYSA-N valinyl-arginine Natural products CC(C)C(N)C(=O)NC(C(O)=O)CCCN=C(N)N IBIDRSSEHFLGSD-UHFFFAOYSA-N 0.000 description 1
- 108010073969 valyllysine Proteins 0.000 description 1
- 238000005406 washing Methods 0.000 description 1
Images
Classifications
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/11—DNA or RNA fragments; Modified forms thereof; Non-coding nucleic acids having a biological activity
- C12N15/113—Non-coding nucleic acids modulating the expression of genes, e.g. antisense oligonucleotides; Antisense DNA or RNA; Triplex- forming oligonucleotides; Catalytic nucleic acids, e.g. ribozymes; Nucleic acids used in co-suppression or gene silencing
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/02—Methods or apparatus for hybridisation; Artificial pollination ; Fertility
-
- 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
- C12N15/64—General methods for preparing the vector, for introducing it into the cell or for selecting the vector-containing host
-
- 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
- C12N2310/00—Structure or type of the nucleic acid
- C12N2310/10—Type of nucleic acid
- C12N2310/20—Type of nucleic acid involving clustered regularly interspaced short palindromic repeats [CRISPRs]
Landscapes
- Health & Medical Sciences (AREA)
- Genetics & Genomics (AREA)
- Life Sciences & Earth Sciences (AREA)
- Engineering & Computer Science (AREA)
- Biomedical Technology (AREA)
- Zoology (AREA)
- General Engineering & Computer Science (AREA)
- Wood Science & Technology (AREA)
- Bioinformatics & Cheminformatics (AREA)
- Chemical & Material Sciences (AREA)
- Organic Chemistry (AREA)
- Biotechnology (AREA)
- Molecular Biology (AREA)
- General Health & Medical Sciences (AREA)
- Biophysics (AREA)
- Plant Pathology (AREA)
- Physics & Mathematics (AREA)
- Microbiology (AREA)
- Biochemistry (AREA)
- Cell Biology (AREA)
- Environmental Sciences (AREA)
- Developmental Biology & Embryology (AREA)
- Botany (AREA)
- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
- Micro-Organisms Or Cultivation Processes Thereof (AREA)
Abstract
本发明公开了一种单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法。本发明首先提供了核苷酸序列为SEQ ID No.1和2所示的靶向OsLG1基因的靶标序列以及SEQ ID No.3所示的靶向OsMTL基因的靶标序列。本发明应用该靶标序列构建得到基因编辑载体。本发明采用该基因编辑载体建立一种结合单倍体育种技术和基因编辑技术改良水稻株型或耐密育种的方法。本发明方法最终的基因编辑体在单倍体诱导过程中产生，不依赖目标材料的遗传转化能力；从单倍体诱导到产生纯合稳定的二倍体基因编辑体，只需两个世代；含有基因编辑载体的父本染色体在单倍体诱导过程中被排除掉，最终得到的基因编辑体不带有转基因载体。
Description
技术领域
本发明涉及改良水稻株型的方法，尤其涉及单倍体介导培育Oslg1突变体表型改良水稻株型或提高产量的方法，属于水稻株型改良的育种领域。
背景技术
水稻是中国主要的粮食作物，占中国口粮消费60％。水稻的充足供给对于保障中国粮食安全意义重大。然而，近年来，人口数量的增长及耕地面积的制约，迫使不得不把提高单位面积的水稻产量作为提高水稻产出的主要技术措施。另一方面，中国乃至世界水稻生产始终面临着日益加剧的病虫害、灾害性天气等逆境威胁，并且依赖化肥和农药的过量施用带动水稻产量增加的效果已趋极限并对中国的农业生态环境造成很大破坏，这也对作物品种的更新速度和效率提出了更高的要求和更严峻的挑战，尤其是耐密新品种的培育。
提高水稻种植密度、增加单位面积的有效穗数是提高水稻单产的主要技术措施。而研究表明，降低植株的茎叶夹角是提高水稻耐密性的关键。较小的茎叶夹角可以最大程度上减少水稻植株之间的相互遮荫，改善田间整体的冠层结构，增强植株间的通风透光性，利于水稻捕获阳光和进行光合作用，利于水稻高产；同时良好的通风透光还将大大增加植株下层的红光/远红光(R/FR)的比例，减少密植避荫反应造成的茎秆徒长、根系变弱、茎秆强度降低等不利影响，利于水稻稳产；此外，研究表明，紧凑的叶夹角还有利于叶片对氮素的同化促进灌浆，直接影响水稻产量。
已有报道显示，OsLG1基因在调控水稻叶夹角及落粒性方面有重要的作用。OsLG1基因突变以后，水稻叶片的叶耳叶舌消失，茎叶夹角显著变小，植株整体上变得紧凑。同时水稻OsLG1的低表达能显著减小稻穗主茎与穗分枝及小花的夹角，从而大大减小水稻的落粒性，并在水稻驯化育种过程中受到选择。
优良基因资源的育种应用，离不开快速、高效育种技术的开发；而传统作物定向改良的育种方法一般需要至少6个以上世代的连续回交和筛选，育种周期长、成本高；并且受遗传累赘效应的影响，性状改良往往不彻底，育种效率低。另一方面，作为当前作物生产领域的核心育种技术，单倍体育种技术可缩短育种周期，但育种的方向较难控制，盲目性大；基因编辑技术可精准地进行目标性状的改良，但所涉及的遗传转化过程受制于材料的遗传背景，且必需的CRISRP载体的去除过程还额外增加育种周期。因此，生产上迫切需要一种能够快速、精准改良作物性状的育种方法，以加快作物育种的进程。
发明内容
本发明的目的之一是提供特异对OsLG1基因进行基因编辑的靶标序列及利用该靶标序列所构建的基因编辑载体；
本发明的目的之二是提供特异对OsMTL基因进行基因编辑的靶标序列及利用该靶标序列所构建的基因编辑载体；
本发明的目的之三是应用所述的基因编辑载体实现两代内对OsLG1 基因进行精准高效的定点编辑培育Oslg1突变体表型二倍体水稻的方法。
本发明的上述目的是通过以下技术方案来实现的：
本发明首先提供了一种特异对OsLG1基因进行基因编辑的靶标DNA序列，其多核苷酸为(a)、(b)、(c)或(d)或所示：
(a)SEQ ID No.1或SEQ ID No.2所示的多核苷酸序列；
(b)与SEQ ID No.1或SEQ ID No.2的互补序列在严谨杂交条件能够进行杂交的多核苷酸；
(c)与SEQ ID No.1或SEQ ID No.2所示的多核苷酸至少有90％或以上同源性的多核苷酸；
(d)在SEQ ID No.1或SEQ ID No.2所示的多核苷酸的基础上进行一个或多个碱基的缺失、取代或插入的多核苷酸突变体，且该多核苷酸突变体仍具有靶向OsLG1基因进而调控水稻叶夹角的功能或活性。
所述的靶标DNA序列能够靶向OsLG1基因并介导Cas9/Cpf1等蛋白对其进行编辑，进而实现调控水稻叶夹角的功能。
利用SEQ ID No.1或SEQ ID No.2的靶标DNA序列所构建的基因编辑载体也应属于本发明的保护范围，这里所指的基因编辑载体中所用的切割核苷酸的蛋白应包括但不限于Cas9、Cpf1及其变体等一切具有核苷酸切割活性的蛋白。
作为本发明的一种具体实施方式，本发明提供了一种所述编辑OsLG1 基因的基因编辑载体的构建方法，包括：(1)将人类中的hSpCas9序列克隆到pCPB载体中构建得到pCPB-Ubi:hSpCas9载体：(2)将所述的靶标DNA 序列引入到sgRNA表达盒中；(3)将引入了靶标DNA序列的sgRNA表达盒通过插入到pCPB-Ubi:hSpCas9的HindIII酶切位点间，得到编辑OsLG1基因的基因编辑载体，其筛选标记是草铵膦抗性。
本发明还提供了特异对OsMTL基因进行基因编辑的靶标DNA序列，其多核苷酸为(a)、(b)、(c)或(d)或所示：
(a)SEQ ID No.3所示的多核苷酸序列；
(b)与SEQ ID No.3的互补序列在严谨杂交条件能够进行杂交的多核苷酸；
(c)与SEQ ID No.3所示的多核苷酸至少有90％或以上同源性的多核苷酸；
(d)在SEQ ID No.3所示的多核苷酸的基础上进行一个或多个碱基的缺失、取代或插入的多核苷酸突变体。
利用SEQ ID No.3的靶标DNA序列所构建的OsMTL基因编辑载体也应属于本发明的保护范围，这里所指的基因编辑载体中所用的切割核苷酸的蛋白应包括但不限于Cas9、Cpf1及其变体等一切具有核苷酸切割活性的蛋白。
作为本发明的一种具体实施方式，本发明提供了一种所述编辑OsMTL 基因的基因编辑载体的构建方法，包括：(1)将人类中的hSpCas9序列克隆到pCPB载体中构建得到pCPB-Ubi:hSpCas9载体：(2)将所述的靶标DNA 序列引入到sgRNA表达盒中；(3)将引入了靶标DNA序列的sgRNA表达盒通过插入到pCPB-Ubi:hSpCas9的HindIII酶切位点间，得到编辑OsMTL基因的基因编辑载体，其筛选标记是潮霉素抗性。
可将本发明提供的靶标DNA序列、该靶标DNA序列的突变体结合“单倍体介导的基因编辑技术”及创制的OsLG1编辑事件等应用于耐密植的水稻新品种，尤其是应用在改良叶夹角和株型、提高水稻产量等方面。
本发明更进一步提供了一种利用单倍体介导的基因编辑技术进行OsLG1 基因编辑改良水稻株型或提高产量的方法，包括：(1)将所述的编辑OsLG1 基因的基因编辑载体以及所述的编辑OsMTL基因的基因编辑载体共转化水稻；(2)在所转化的后代中分离挑选出两个OsMTL基因被编辑突变、携带有编辑OsLG1基因的基因编辑载体、但不携带编辑OsMTL基因的基因编辑载体的T2株系；(3)用这两个T2株系的花粉与需要进行基因编辑的材料进行杂交诱导单倍体，获得OsLG1基因发生编辑突变的单倍体植株；(4)将该单倍体进行染色体加倍得到带有编辑后的OsLG1基因、能稳定遗传、叶夹角变小、不含有转基因载体的二倍体水稻材料。
作为一种优选的具体实施方式，步骤(2)中两个T2株系的筛分方法包括：(a)将共转化后的T0植株用潮霉素和草铵膦除草剂进行筛选，挑选出双阳性的转基因水稻进行自交，得到T1代幼苗；(b)用潮霉素和草铵膦除草剂对T1代幼苗进行筛选得到具有草铵膦抗性、但不具有潮霉素抗性的单株；(c)对单株进行OsMTL基因的测序，找到两株OsMTL基因发生纯合突变的单株；(d)对这两个单株进行自交得到两个T2代株系。
作为一种优选的具体实施方式，步骤(2)中所述的两个OsMTL基因被编辑突变后得到的两个突变体的核苷酸序列分别为SEQ ID No.8和SEQ ID No.9所示；步骤(3)中所述的OsLG1基因发生编辑突变后得到的突变体的核苷酸序列为SEQ ID No.10所示。
本发明中所述OsLG1基因，其编码区的核苷酸序列为SEQ ID No.4所示，其编码蛋白如SEQ ID No.5所示。
本发明中所述OsMTL基因，其编码区的核苷酸序列为SEQ ID No.6 所示，其编码蛋白如SEQ ID No.7所示。
进一步的，含有所述SEQ ID No.1所述的DNA序列、SEQ ID No.2所示的DNA序列或SEQ ID No.3所示的DNA序列及其突变体的表达盒、含有该表达盒的重组植物表达载体、转基因细胞系和宿主菌均属于本发明的保护范围。
所述的重组植物表达载体是将所述表达盒与质粒或表达载体所构建得到的重组植物表达载体并且能够将所述表达盒转入植物宿主细胞、组织或器官中。
本发明的DNA序列或其突变体可以用于制备转基因植物。譬如，通过农杆菌介导、基因枪法等方法将含有所述DNA序列或其突变体的重组植物表达载体导入植物细胞、组织或器官中，再将该转化的植物细胞、组织或器官培育成植株，获得转基因植物。用于构建所述植物表达载体的出发载体可为任意一种用于农杆菌转化植物的双元载体或可用于植物微弹轰击的载体等。
为了实施本发明，制备和使用植物表达载体和宿主细胞的常规组合物和方法为本领域技术人员所熟知，具体方法可以参考例如Sambrook等。
所述重组植物表达载体还可含有用于选择转化细胞的选择性标记基因。选择性标记基因用于选择经转化的细胞或组织。所述的标记基因包括：编码抗生素抗性的基因以及赋予除草化合物抗性的基因等。此外，所述的标记基因还包括表型标记，例如β-半乳糖苷酶和荧光蛋白等。
本发明中所述的转化方案以及将所述多核苷酸或多肽引入植物的方案可视用于转化的植物(单子叶植物或双子叶植物)或植物细胞的类型而变化。将所述多核苷酸或多肽引入植物细胞的合适方法包括：显微注射、电穿孔、农杆菌介导的转化、直接基因转移以及高速弹道轰击等。在特定的实施方案中，可利用多种瞬时转化法将本发明的表达盒提供给植物。利用常规方法可使已转化的细胞再生稳定转化植株(McCormick et al.Plant CellReports.1986. 5:81-84)。
本发明可用于转化任何植物种类，包括但不限于：单子叶植物或双子叶植物，优选为水稻。
基于OsLG1基因在调控水稻叶夹角及落粒性方面的重要作用结合单倍体育种技术和基因编辑技术的特点，本发明采用特异对OsLG1基因进行基因编辑的靶标DNA序列巧妙结合单倍体育种技术和基因编辑技术来改良水稻株型并用于水稻耐密育种。本发明方法最终的基因编辑体在单倍体诱导的过程中产生，不依赖目标材料的遗传转化能力；从单倍体诱导到最终产生纯合稳定的二倍体基因编辑体，只需两个世代(一次诱导、一次加倍)，快捷方便、节省成本；含有基因编辑载体的父本染色体在单倍体诱导过程中被排除掉，因此该发明最终得到的基因编辑体不带有转基因载体。
本发明所涉及到的术语定义
除非另外定义，否则本文所用的所有技术及科学术语都具有与本发明所属领域的普通技术人员通常所了解相同的含义。虽然在本发明的实践或测试中可使用与本文所述者类似或等效的任何方法、装置和材料，但现在描述优选方法、装置和材料。
在本发明的上下文中，术语“突变体”是含有变化的DNA序列，在所述 DNA序列中，优选在基本保持DNA序列的同时缺失、添加和/或替代原始序列的一个或多个核苷酸。例如，可以从DNA序列5’或3’末端缺失一个或多个碱基对以产生“截短的”DNA序列；也可以在DNA序列内部插入、缺失或替代一个或多个碱基对。可以通过例如标准DNA诱变技术或通过化学合成变体DNA序列或其部分而产生变体DNA序列。突变体多核苷酸还包括合成来源的多核苷酸，例如采用定点诱变所得到的突变体，或者是通过重组的方法(例如DNA改组)所得到的突变体，或者是通过自然选择所得到的突变体。
术语“多核苷酸”或“核苷酸”意指单股或双股形式的脱氧核糖核苷酸、脱氧核糖核苷、核糖核苷或核糖核苷酸及其聚合物。除非特定限制，否则所述术语涵盖含有天然核苷酸的已知类似物的核酸，所述类似物具有类似于参考核酸的结合特性并以类似于天然产生的核苷酸的方式进行代谢。除非另外特定限制，否则所述术语也意指寡核苷酸类似物，其包括PNA(肽核酸)、在反义技术中所用的DNA类似物(硫代磷酸酯、磷酰胺酸酯等等)。除非另外指定，否则特定核酸序列也隐含地涵盖其保守修饰的变异体(包括(但不限于) 简并密码子取代)和互补序列以及明确指定的序列。特定而言，可通过产生其中一个或一个以上所选(或所有)密码子的第3位经混合碱基和/或脱氧肌苷残基取代的序列来实现简并密码子取代(Batzer等人,Nucleic Acid Res. 19:5081(1991)；Ohtsuka等人,J.Biol.Chem.260:2605-2608(1985)；和Cassol 等人,(1992)；Rossolini等人,Mol Cell.Probes 8:91-98(1994))。
术语“同源性”指多核苷酸序列之间在百分比核苷酸位置同一性(即序列相似性或同一性)方面的相似性或百分同一性的水平。此处所用的术语同源性也指不同多核苷酸分子之间相似的功能特性的概念，例如具有相似功能的启动子可能具有同源的顺式元件。当多核苷酸分子在特定条件下特异性地杂交以形成双链体分子时，它们是同源的。在这些条件下(称为严谨杂交条件)一个多核苷酸分子可以用作鉴定共有同源性的另一个多核苷酸分子的探针或引物。
本发明中所述“严谨杂交条件”意指在所属领域中已知的低离子强度和高温的条件。通常，在严谨条件下，探针与其靶序列杂交的可检测程度比与其它序列杂交的可检测程度更高(例如超过本底至少2倍。严谨杂交条件是序列依赖性的，在不同的环境条件下将会不同，较长的序列在较高温度下特异性杂交。通过控制杂交的严谨性或洗涤条件可鉴定与探针100％互补的靶序列。对于核酸杂交的详尽指导可参考有关文献(Tijssen,Techniquesin Biochemistry and Molecular Biology-Hybridization with Nucleic Probes, "Overview of principles of hybridization and the strategy of nucleic acidassays.1993)。更具体的，所述严谨条件通常被选择为低于特异序列在规定离子强度pH下的热熔点(Tm)约5-10℃。Tm为在平衡状态下50％与目标互补的探针杂交到目标序列时所处的温度(在指定离子强度、pH和核酸浓度下) (因为目标序列过量存在，所以在Tm下在平衡状态下50％的探针被占据)。严谨条件可为以下条件：其中在pH 7.0到8.3下盐浓度低于约1.0M钠离子浓度，通常为约0.01到1.0M钠离子浓度(或其它盐)，并且温度对于短探针 (包括(但不限于)10到50个核苷酸)而言为至少约30℃，而对于长探针 (包括(但不限于)大于50个核苷酸)而言为至少约60℃。严谨条件也可通过加入诸如甲酰胺的去稳定剂来实现。对于选择性或特异性杂交而言，正信号可为至少两倍的背景杂交，视情况为10倍背景杂交。例示性严谨杂交条件可如下：50％甲酰胺，5×SSC和1％SDS，在42℃下培养；或5×SSC，1％SDS，在65℃下培养，在0.2×SSC中洗涤和在65℃下于0.1％SDS中洗涤。所述洗涤可进行5、15、30、60、120分钟或更长时间。
本发明中所述的“多个”通常意味着2-8个，优选为2-4个；所述的“替换”是指分别用不同的氨基酸残基取代一个或多个氨基酸残基；所述的“缺失”是指氨基酸残基数量的减少，也即是分别缺少其中的一个或多个氨基酸残基；所述的“插入”是指氨基酸残基序列的改变，相对天然分子而言，所述改变导致添加一个或多个氨基酸残基。
术语“编码序列”：转录成RNA的核酸序列。
术语“转化”：将异源性DNA序列引入到宿主细胞或有机体的方法。
术语“表达”：内源性基因或转基因在植物细胞中的转录和/或翻译。
术语“重组宿主细胞株”或“宿主细胞”意指包含本发明多核苷酸的细胞，而不管使用何种方法进行插入以产生重组宿主细胞，例如直接摄取、转导、f配对或所属领域中已知的其它方法。外源性多核苷酸可保持为例如质粒的非整合载体或者可整合入宿主基因组中。宿主细胞可为原核细胞或真核细胞，宿主细胞还可为单子叶或双子叶植物细胞。
附图说明
图1为所构建的靶向OsMTL基因的基因编辑载体；该载体中所用的筛选标记为潮霉素抗性基因。
图2为所构建的靶向OsLG1基因的基因编辑载体；该载体中所用的筛选标记为草铵膦抗性基因。
图3为本发明中利用靶向OsLG1的“单倍体介导的基因编辑系统”改良水稻叶夹角的流程图。
图4为本发明所创制的两个OsMTL基因突变体的测序情况。
图5A：所创制的野生型材料(Wild-type)、正常水稻单倍体 (WT-Haploid)和OsLG1基因发生编辑的单倍体(Oslg1-Haploid)的叶夹角情况；图5B：为野生型材料、正常水稻单倍体和OsLG1基因发生编辑的单倍体的测序情况。
图6所创制的野生型材料(Wild-type)、正常水稻单倍体(WT-Haploid) 和OsLG1基因发生编辑的单倍体(Oslg1-Haploid)的流式细胞仪倍性检测情况。
图7为所用野生型及所创制的Oslg1改良材料的植株、叶夹角和稻穗照片。
具体实施方式
以下结合具体实施例来进一步描述本发明，本发明的优点和特点将会随着描述而更为清楚。但这些实施例仅是范例性的，并不对本发明的范围构成任何限制。本领域技术人员应该理解的是，在不偏离本发明的精神和范围下可以对本发明的细节和形式进行修改或替换，但这些修改和替换均落入本发明的保护范围内。
以下实施例中所用的水稻材料可从“中国作物种质信息网”得到相关信息和申请获取对应的种子。
实施例1、OsMTL-CRISPR/CAS9和OsLG1-CRISPR/CAS9基因编辑载体的构建和转化
首先，利用SnapGene Viewer软件和OsMTL与OsLG1的基因序列初步查找其靶标编辑序列；所获得的序列再与水稻“日本晴”(Japonica) 的其他基因组序列进行比对，以去除多拷贝和与其他基因组区域高度相似的序列；根据最终确定的靶标序列进一步设计sgRNA，并通过RNA Folding Form(http://unafold.rna.albany.edu/？q＝mfold/RNA-Folding-Form2.3)系统来进行二级结构的预测；最终选取设计了最优的靶标序列用于编辑OsMTL 与OsLG1基因；其中OsMTL基因设计了一个靶标序列(SEQ ID No.3)， OsLG1基因设计了两个靶标序列(分别为SEQ ID No.1和SEQ ID No.2所示)。之后，将人类中的hSpCas9序列用商业化的
PCR Cloning Kit试剂盒克隆到pCPB载体中，构建成为pCPB-Ubi:hSpCas9载体。之后用重叠PCR的方法将之前获得的靶标序列引入到sgRNA表达盒中。接着，将sgRNA表达盒通过
HD Cloning Kit试剂盒插入到 pCPB-Ubi:hSpCas9的HindIII酶切位点间。其中OsMTL-CRISPR/CAS9所用的筛选标记是潮霉素抗性，OsLG1-CRISPR/CAS9所用的筛选标记是草铵膦抗性(图1和图2)，最终构建成的CRISPR/Cas9基因编辑编辑载体(图1和图2)经过PCR测序验证无误后用于后续的遗传转化。两个载体同时转化同一批次水稻愈伤组织；该载体的遗传转化通过常规的农杆菌介导的方法完成，所转化的受体材料为水稻“日本晴”。
实施例2、创制OsMTL基因被编辑、携带OsLG1-CRISPR/CAS9载体、但不携带OsMTL-CRISPR/CAS9载体的水稻单倍体诱导系
OsMTL-CRISPR/CAS9和OsLG1-CRISPR/CAS9基因编辑载体共转化水稻，得到大约40株T0代植株；将这40株T0植株用潮霉素和草铵膦除草剂进行筛选，挑选出6株双阳性的转基因水稻；对这些双阳性植株进行自交，得的120株T1代幼苗；用潮霉素和草铵膦除草剂对这些T1代幼苗进行筛选，找到了21株具有草铵膦抗性、但不具有潮霉素抗性的单株(携带有OsLG1-CRISPR/CAS9基因编辑载体，但不携带OsMTL-CRISPR/CAS9基因编辑载体)；进而对这些单株进行OsMTL基因的测序，找到两株OsMTL 基因发生纯合突变的单株(图4)，这两个单株的基因型为OsMTL基因发生纯合突变、携带有OsLG1-CRISPR/CAS9基因编辑载体，但不携带 OsMTL-CRISPR/CAS9基因编辑载体。对他们进行自交得到了两个T2代株系。这2个T2株系因OsMTL基因突变所以具有单倍体诱导的能力，同时因携带有OsLG1-CRISPR/CAS9基因编辑载体所以具有编辑OsLG1基因的能力。
实施例3、利用“OsMTL基因突变、携带OsLG1-CRISPR/CAS9载体、但不携带OsMTL-CRISPR/CAS9载体的水稻单倍体诱导系”进行水稻OsLG1 编辑改良
利用所创制的两个携带OsLG1-CRISPR/CAS9载体的水稻单倍体诱导系的花粉与水稻品种“中花11”进行杂交，之后所得种子播种至苗床后，共得到609株杂交后代；根据生长势，去除长势壮的双倍体，得到约23株准单倍体幼苗；其中得到一株具有OsLG1突变表型的准单倍体幼苗(图5)。测序分析证明该单株的OsLG1基因在第二个靶标位点处发生了一个碱基的插入，造成了OsLG1基因的移码突变。利用流式细胞仪对正常的野生型、正常的单倍体及具有Oslg1突变体表型的植株进行倍性检测证明了该单株确实是单倍体(图6)。利用秋水仙素处理该单株的叶心，成功实现了该单株的染色体加倍，得到了具有Oslg1突变体表型的正常二倍体水稻(图 7)。整个过程，从单倍体诱导到产生Oslg1突变体表型的正常二倍体水稻，只用了2个世代。
SEQUENCE LISTING
<110> 华南农业大学 中国农业科学院生物技术研究所
<120> 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法
<130> BJ-2002-200801A
<160> 10
<170> PatentIn version 3.5
<210> 1
<211> 22
<212> DNA
<213> Artifical sequence
<400> 1
gaactggagg aggtcatgtg gg 22
<210> 2
<211> 22
<212> DNA
<213> Artifical sequence
<400> 2
ggtttcgaga cacagctact gg 22
<210> 3
<211> 26
<212> DNA
<213> Artifical sequence
<400> 3
gccgggcacc tcgacgtacc tgccgg 26
<210> 4
<211> 1251
<212> DNA
<213> Artifical sequence
<400> 4
atgatgaacg ttccatccgc cgctgccgcg agctcctgcg atgatttcgg ctacaacgcc 60
accccgccgc cgccgccgtc gcttctccca atcatggacc aggacggcgg cggcggtagc 120
atccagaggg atcaccacca acaccacaac caccagcagc tcggctacaa cctggagccg 180
agctccctag ctctgcttcc cccttccaac gccgccgccg ccgcagcaca ccacgccacc 240
atcgcccacg cctccccaca tgacctcctc cagttctacc cgacctcgca ctacctcgcc 300
gctgccggcg gcgccggtgg cggaggcaac ccctacagcc acttcacggc ggcggcggcg 360
gccgggagca ccttccagtc gtactaccag cagccgccgc aggacgcgcc ggagtactac 420
ttcccgacgc tggtcagctc cgccgaggag aacatggcca gcttcgccgc cacccagctc 480
ggcctcaacc tcggctaccg gacctacttc ccgccgagag gcggctacac ctacggccac 540
cacccgccgc ggtgccaggc cgagggctgc aaggccgacc tctccagcgc caagcgctac 600
caccgccgcc acaaggtgtg cgagcaccac tccaaggcgc ccgtcgtcgt caccgccggc 660
ggcctccacc agagattctg ccagcagtgc agcagattcc atcttcttga tgagttcgac 720
gatgccaaga agagctgcag gaagcgactc gccgaccaca accgccggcg gaggaagtcg 780
aagccgtccg acggcgagca ttctggtgaa aagagaaggg cgcaggcgaa taaatcggca 840
gctactaaag acaaagcagg aagtagcagc aagaacgcag gcattggaga cggtttcgag 900
acacagctac tggggggtgc acacatgtcc aaagatcaag accaagccat ggatctggga 960
gaggtggtga aagaagctgt agatcccaaa ggtaaggcat cgatgcagca gcagcagcag 1020
caagcacatc atgggattca tcagcagagc caccagcagc atggcttccc tttcccttcg 1080
tcgtctggct cgtgcttatt ccctcagagc caaggagctg tctcgagcac tgacacatca 1140
aatatagctc aagtgcaaga accaagctta gccttccatc agcagcatca ccaacacagc 1200
aacatccttc agcttggaca ggcgatgttt gatctcgact tcgatcacta g 1251
<210> 5
<211> 416
<212> PRT
<213> Artifical sequence
<400> 5
Met Met Asn Val Pro Ser Ala Ala Ala Ala Ser Ser Cys Asp Asp Phe
1 5 10 15
Gly Tyr Asn Ala Thr Pro Pro Pro Pro Pro Ser Leu Leu Pro Ile Met
20 25 30
Asp Gln Asp Gly Gly Gly Gly Ser Ile Gln Arg Asp His His Gln His
35 40 45
His Asn His Gln Gln Leu Gly Tyr Asn Leu Glu Pro Ser Ser Leu Ala
50 55 60
Leu Leu Pro Pro Ser Asn Ala Ala Ala Ala Ala Ala His His Ala Thr
65 70 75 80
Ile Ala His Ala Ser Pro His Asp Leu Leu Gln Phe Tyr Pro Thr Ser
85 90 95
His Tyr Leu Ala Ala Ala Gly Gly Ala Gly Gly Gly Gly Asn Pro Tyr
100 105 110
Ser His Phe Thr Ala Ala Ala Ala Ala Gly Ser Thr Phe Gln Ser Tyr
115 120 125
Tyr Gln Gln Pro Pro Gln Asp Ala Pro Glu Tyr Tyr Phe Pro Thr Leu
130 135 140
Val Ser Ser Ala Glu Glu Asn Met Ala Ser Phe Ala Ala Thr Gln Leu
145 150 155 160
Gly Leu Asn Leu Gly Tyr Arg Thr Tyr Phe Pro Pro Arg Gly Gly Tyr
165 170 175
Thr Tyr Gly His His Pro Pro Arg Cys Gln Ala Glu Gly Cys Lys Ala
180 185 190
Asp Leu Ser Ser Ala Lys Arg Tyr His Arg Arg His Lys Val Cys Glu
195 200 205
His His Ser Lys Ala Pro Val Val Val Thr Ala Gly Gly Leu His Gln
210 215 220
Arg Phe Cys Gln Gln Cys Ser Arg Phe His Leu Leu Asp Glu Phe Asp
225 230 235 240
Asp Ala Lys Lys Ser Cys Arg Lys Arg Leu Ala Asp His Asn Arg Arg
245 250 255
Arg Arg Lys Ser Lys Pro Ser Asp Gly Glu His Ser Gly Glu Lys Arg
260 265 270
Arg Ala Gln Ala Asn Lys Ser Ala Ala Thr Lys Asp Lys Ala Gly Ser
275 280 285
Ser Ser Lys Asn Ala Gly Ile Gly Asp Gly Phe Glu Thr Gln Leu Leu
290 295 300
Gly Gly Ala His Met Ser Lys Asp Gln Asp Gln Ala Met Asp Leu Gly
305 310 315 320
Glu Val Val Lys Glu Ala Val Asp Pro Lys Gly Lys Ala Ser Met Gln
325 330 335
Gln Gln Gln Gln Gln Ala His His Gly Ile His Gln Gln Ser His Gln
340 345 350
Gln His Gly Phe Pro Phe Pro Ser Ser Ser Gly Ser Cys Leu Phe Pro
355 360 365
Gln Ser Gln Gly Ala Val Ser Ser Thr Asp Thr Ser Asn Ile Ala Gln
370 375 380
Val Gln Glu Pro Ser Leu Ala Phe His Gln Gln His His Gln His Ser
385 390 395 400
Asn Ile Leu Gln Leu Gly Gln Ala Met Phe Asp Leu Asp Phe Asp His
405 410 415
<210> 6
<211> 1299
<212> DNA
<213> Artifical sequence
<400> 6
atggcggcga gctactcgtg ccggcggaca tgcgaggcgt gcagcacgag ggcgatggcc 60
gggtgcgtgg tgggcgagcc ggcgtcggcg ccggggcagc gggtgacgtt gctggcgatc 120
gacggcggcg gcatcagggg cctcatcccg ggcaccatcc tcgccttcct cgaggccagg 180
ctgcaggagc tggatggccc cgacgcgcgc ctcgccgatt acttcgactg catcgccggg 240
accagcaccg gcggcctcat caccgccatg ctcgccgcgc ccggcgacca cggccgcccg 300
ctcttcgccg ccagcgacat caaccgcttc tacctcgaca acggcccact catcttccca 360
caaaagaggt gcggcatggc ggcggccatg gcggcgctga cgaggccgag gtacaacggc 420
aagtacctgc aggggaagat caggaagatg ctgggcgaga cgagggtgcg cgacacgctg 480
acgaacgtcg tcatccccac gttcgacgtc aggctgctcc agccaaccat cttctccaca 540
tacgacgcga agagcatgcc gctcaagaac gcgctcctct ccgacatctg catcagcaca 600
tccgcggcgc cgacctacct ccccgcgcac tgcttccaga ccaccgacga cgccaccggc 660
aaggtccgcg agttcgacct catcgacggc ggcgtcgccg ccaacaaccc gacgatggtg 720
gccatgacgc agatcaccaa gaagataatg gtgaaggaca aggaggagct gtacccggta 780
aagccgtcgg actgcggtaa gttcctggtg ctgtccgtgg gcaccgggtc gacgtcggac 840
caggggatgt acacggcgag gcagtgctcg cggtggggga tcgtccggtg gctgcgcaac 900
aaggggatgg cgcccatcat cgacatcttc atggcggcca gctccgacct cgtcgacatc 960
cacgccgccg tcatgttcca gtcgctgcac agcgacggcg actacctccg catccaggac 1020
aacacgctcc acggcgacgc cgccacggtg gacgccgcca ccagggacaa catgcgggcg 1080
ctcgtcggga tcggcgagcg gatgctggcg cagcgggtgt cgagggtcaa cgtcgagacc 1140
ggcaggtacg tcgaggtgcc cggcgccggc agcaacgccg acgcgctgag gggcttcgcc 1200
aggcagctct ccgaggagag gagggcgagg ctaggtcggc gaaacgcctg cggcggcggc 1260
ggcgaaggag agcccagcgg cgtggcgtgc aagcgttag 1299
<210> 7
<211> 432
<212> PRT
<213> Artifical sequence
<400> 7
Met Ala Ala Ser Tyr Ser Cys Arg Arg Thr Cys Glu Ala Cys Ser Thr
1 5 10 15
Arg Ala Met Ala Gly Cys Val Val Gly Glu Pro Ala Ser Ala Pro Gly
20 25 30
Gln Arg Val Thr Leu Leu Ala Ile Asp Gly Gly Gly Ile Arg Gly Leu
35 40 45
Ile Pro Gly Thr Ile Leu Ala Phe Leu Glu Ala Arg Leu Gln Glu Leu
50 55 60
Asp Gly Pro Asp Ala Arg Leu Ala Asp Tyr Phe Asp Cys Ile Ala Gly
65 70 75 80
Thr Ser Thr Gly Gly Leu Ile Thr Ala Met Leu Ala Ala Pro Gly Asp
85 90 95
His Gly Arg Pro Leu Phe Ala Ala Ser Asp Ile Asn Arg Phe Tyr Leu
100 105 110
Asp Asn Gly Pro Leu Ile Phe Pro Gln Lys Arg Cys Gly Met Ala Ala
115 120 125
Ala Met Ala Ala Leu Thr Arg Pro Arg Tyr Asn Gly Lys Tyr Leu Gln
130 135 140
Gly Lys Ile Arg Lys Met Leu Gly Glu Thr Arg Val Arg Asp Thr Leu
145 150 155 160
Thr Asn Val Val Ile Pro Thr Phe Asp Val Arg Leu Leu Gln Pro Thr
165 170 175
Ile Phe Ser Thr Tyr Asp Ala Lys Ser Met Pro Leu Lys Asn Ala Leu
180 185 190
Leu Ser Asp Ile Cys Ile Ser Thr Ser Ala Ala Pro Thr Tyr Leu Pro
195 200 205
Ala His Cys Phe Gln Thr Thr Asp Asp Ala Thr Gly Lys Val Arg Glu
210 215 220
Phe Asp Leu Ile Asp Gly Gly Val Ala Ala Asn Asn Pro Thr Met Val
225 230 235 240
Ala Met Thr Gln Ile Thr Lys Lys Ile Met Val Lys Asp Lys Glu Glu
245 250 255
Leu Tyr Pro Val Lys Pro Ser Asp Cys Gly Lys Phe Leu Val Leu Ser
260 265 270
Val Gly Thr Gly Ser Thr Ser Asp Gln Gly Met Tyr Thr Ala Arg Gln
275 280 285
Cys Ser Arg Trp Gly Ile Val Arg Trp Leu Arg Asn Lys Gly Met Ala
290 295 300
Pro Ile Ile Asp Ile Phe Met Ala Ala Ser Ser Asp Leu Val Asp Ile
305 310 315 320
His Ala Ala Val Met Phe Gln Ser Leu His Ser Asp Gly Asp Tyr Leu
325 330 335
Arg Ile Gln Asp Asn Thr Leu His Gly Asp Ala Ala Thr Val Asp Ala
340 345 350
Ala Thr Arg Asp Asn Met Arg Ala Leu Val Gly Ile Gly Glu Arg Met
355 360 365
Leu Ala Gln Arg Val Ser Arg Val Asn Val Glu Thr Gly Arg Tyr Val
370 375 380
Glu Val Pro Gly Ala Gly Ser Asn Ala Asp Ala Leu Arg Gly Phe Ala
385 390 395 400
Arg Gln Leu Ser Glu Glu Arg Arg Ala Arg Leu Gly Arg Arg Asn Ala
405 410 415
Cys Gly Gly Gly Gly Glu Gly Glu Pro Ser Gly Val Ala Cys Lys Arg
420 425 430
<210> 8
<211> 1277
<212> DNA
<213> Artifical sequence
<400> 8
atggcggcga gctactcgtg ccggcggaca tgcgaggcgt gcagcacgag ggcgatggcc 60
gggtgcgtgg tgggcgagcc ggcgtcggcg ccggggcagc gggtgacgtt gctggcgatc 120
gacggcggcg gcatcagggg cctcatcccg ggcaccatcc tcgccttcct cgaggccagg 180
ctgcaggagc tggatggccc cgacgcgcgc ctcgccgatt acttcgactg catcgccggg 240
accagcaccg gcggcctcat caccgccatg ctcgccgcgc ccggcgacca cggccgcccg 300
ctcttcgccg ccagcgacat caaccgcttc tacctcgaca acggcccact catcttccca 360
caaaagaggt gcggcatggc ggcggccatg gcggcgctga cgaggccgag gtacaacggc 420
aagtacctgc aggggaagat caggaagatg ctgggcgaga cgagggtgcg cgacacgctg 480
acgaacgtcg tcatccccac gttcgacgtc aggctgctcc agccaaccat cttctccaca 540
tacgacgcga agagcatgcc gctcaagaac gcgctcctct ccgacatctg catcagcaca 600
tccgcggcgc cgacctacct ccccgcgcac tgcttccaga ccaccgacga cgccaccggc 660
aaggtccgcg agttcgacct catcgacggc ggcgtcgccg ccaacaaccc gacgatggtg 720
gccatgacgc agatcaccaa gaagataatg gtgaaggaca aggaggagct gtacccggta 780
aagccgtcgg actgcggtaa gttcctggtg ctgtccgtgg gcaccgggtc gacgtcggac 840
caggggatgt acacggcgag gcagtgctcg cggtggggga tcgtccggtg gctgcgcaac 900
aaggggatgg cgcccatcat cgacatcttc atggcggcca gctccgacct cgtcgacatc 960
cacgccgccg tcatgttcca gtcgctgcac agcgacggcg actacctccg catccaggac 1020
aacacgctcc acggcgacgc cgccacggtg gacgccgcca ccagggacaa catgcgggcg 1080
ctcgtcggga tcggcgagcg gatgctggcg cagcgggtgt cgagggtcaa cgtcgagacg 1140
gcgccggcag caacgccgac gcgctgaggg gcttcgccag gcagctctcc gaggagagga 1200
gggcgaggct aggtcggcga aacgcctgcg gcggcggcgg cgaaggagag cccagcggcg 1260
tggcgtgcaa gcgttag 1277
<210> 9
<211> 1298
<212> DNA
<213> Artifical sequence
<400> 9
atggcggcga gctactcgtg ccggcggaca tgcgaggcgt gcagcacgag ggcgatggcc 60
gggtgcgtgg tgggcgagcc ggcgtcggcg ccggggcagc gggtgacgtt gctggcgatc 120
gacggcggcg gcatcagggg cctcatcccg ggcaccatcc tcgccttcct cgaggccagg 180
ctgcaggagc tggatggccc cgacgcgcgc ctcgccgatt acttcgactg catcgccggg 240
accagcaccg gcggcctcat caccgccatg ctcgccgcgc ccggcgacca cggccgcccg 300
ctcttcgccg ccagcgacat caaccgcttc tacctcgaca acggcccact catcttccca 360
caaaagaggt gcggcatggc ggcggccatg gcggcgctga cgaggccgag gtacaacggc 420
aagtacctgc aggggaagat caggaagatg ctgggcgaga cgagggtgcg cgacacgctg 480
acgaacgtcg tcatccccac gttcgacgtc aggctgctcc agccaaccat cttctccaca 540
tacgacgcga agagcatgcc gctcaagaac gcgctcctct ccgacatctg catcagcaca 600
tccgcggcgc cgacctacct ccccgcgcac tgcttccaga ccaccgacga cgccaccggc 660
aaggtccgcg agttcgacct catcgacggc ggcgtcgccg ccaacaaccc gacgatggtg 720
gccatgacgc agatcaccaa gaagataatg gtgaaggaca aggaggagct gtacccggta 780
aagccgtcgg actgcggtaa gttcctggtg ctgtccgtgg gcaccgggtc gacgtcggac 840
caggggatgt acacggcgag gcagtgctcg cggtggggga tcgtccggtg gctgcgcaac 900
aaggggatgg cgcccatcat cgacatcttc atggcggcca gctccgacct cgtcgacatc 960
cacgccgccg tcatgttcca gtcgctgcac agcgacggcg actacctccg catccaggac 1020
aacacgctcc acggcgacgc cgccacggtg gacgccgcca ccagggacaa catgcgggcg 1080
ctcgtcggga tcggcgagcg gatgctggcg cagcgggtgt cgagggtcaa cgtcgagacc 1140
ggcaggtact cgaggtgccc ggcgccggca gcaacgccga cgcgctgagg ggcttcgcca 1200
ggcagctctc cgaggagagg agggcgaggc taggtcggcg aaacgcctgc ggcggcggcg 1260
gcgaaggaga gcccagcggc gtggcgtgca agcgttag 1298
<210> 10
<211> 1252
<212> DNA
<213> Artifical sequence
<400> 10
atgatgaacg ttccatccgc cgctgccgcg agctcctgcg atgatttcgg ctacaacgcc 60
accccgccgc cgccgccgtc gcttctccca atcatggacc aggacggcgg cggcggtagc 120
atccagaggg atcaccacca acaccacaac caccagcagc tcggctacaa cctggagccg 180
agctccctag ctctgcttcc cccttccaac gccgccgccg ccgcagcaca ccacgccacc 240
atcgcccacg cctccccaca tgacctcctc cagttctacc cgacctcgca ctacctcgcc 300
gctgccggcg gcgccggtgg cggaggcaac ccctacagcc acttcacggc ggcggcggcg 360
gccgggagca ccttccagtc gtactaccag cagccgccgc aggacgcgcc ggagtactac 420
ttcccgacgc tggtcagctc cgccgaggag aacatggcca gcttcgccgc cacccagctc 480
ggcctcaacc tcggctaccg gacctacttc ccgccgagag gcggctacac ctacggccac 540
cacccgccgc ggtgccaggc cgagggctgc aaggccgacc tctccagcgc caagcgctac 600
caccgccgcc acaaggtgtg cgagcaccac tccaaggcgc ccgtcgtcgt caccgccggc 660
ggcctccacc agagattctg ccagcagtgc agcagattcc atcttcttga tgagttcgac 720
gatgccaaga agagctgcag gaagcgactc gccgaccaca accgccggcg gaggaagtcg 780
aagccgtccg acggcgagca ttctggtgaa aagagaaggg cgcaggcgaa taaatcggca 840
gctactaaag acaaagcagg aagtagcagc aagaacgcag gcattggaga cggtttcgag 900
acacagcata ctggggggtg cacacatgtc caaagatcaa gaccaagcca tggatctggg 960
agaggtggtg aaagaagctg tagatcccaa aggtaaggca tcgatgcagc agcagcagca 1020
gcaagcacat catgggattc atcagcagag ccaccagcag catggcttcc ctttcccttc 1080
gtcgtctggc tcgtgcttat tccctcagag ccaaggagct gtctcgagca ctgacacatc 1140
aaatatagct caagtgcaag aaccaagctt agccttccat cagcagcatc accaacacag 1200
caacatcctt cagcttggac aggcgatgtt tgatctcgac ttcgatcact ag 1252
Claims (10)
1.靶向OsLG1基因的靶标DNA序列，其特征在于，其多核苷酸为(a)、(b)、(c)或(d)或所示：
(a)SEQ ID No.1或SEQ ID No.2所示的多核苷酸序列；
(b)与SEQ ID No.1或SEQ ID No.2的互补序列在严谨杂交条件能够进行杂交的多核苷酸；
(c)与SEQ ID No.1或SEQ ID No.2所示的多核苷酸至少有90％或以上同源性的多核苷酸；
(d)在SEQ ID No.1或SEQ ID No.2所示的多核苷酸的基础上进行一个或多个碱基的缺失、取代或插入的多核苷酸突变体，且该多核苷酸突变体仍具有靶向OsLG1基因进而调控水稻叶夹角的功能或活性。
2.含有权利要求1所述靶标DNA序列的编辑OsLG1基因的基因编辑载体。
3.权利要求2所述编辑OsLG1基因的基因编辑载体的构建方法，其特征在于，包括：(1)将人类中的hSpCas9序列克隆到pCPB载体中构建得到pCPB-Ubi:hSpCas9载体：(2)将权利要求1所述的靶标DNA序列引入到sgRNA表达盒中；(3)将引入了靶标DNA序列的sgRNA表达盒插入到pCPB-Ubi:hSpCas9的HindIII酶切位点间得到编辑OsLG1基因的基因编辑载体；该基因编辑载体的筛选标记是草铵膦抗性。
4.一种靶向OsMTL基因的靶标DNA序列，其特征在于，其多核苷酸为(a)、(b)、(c)或(d)或所示：
(a)SEQ ID No.3所示的多核苷酸序列；
(b)与SEQ ID No.3的互补序列在严谨杂交条件能够进行杂交的多核苷酸；
(c)与SEQ ID No.3所示的多核苷酸至少有90％或以上同源性的多核苷酸；
(d)在SEQ ID No.3所示的多核苷酸的基础上进行一个或多个碱基的缺失、取代或插入的多核苷酸突变体。
5.含有权利要求4所述靶标DNA序列的编辑OsMTL基因的基因编辑载体。
6.权利要求5所述编辑OsMTL基因的基因编辑载体的构建方法，其特征在于，包括：(1)将人类中的hSpCas9序列克隆到pCPB载体中构建得到pCPB-Ubi:hSpCas9载体：(2)将权利要求4所述的靶标DNA序列引入到sgRNA表达盒中；(3)将引入了靶标DNA序列的sgRNA表达盒插入到pCPB-Ubi:hSpCas9的HindIII酶切位点间得到编辑OsMTL基因的基因编辑载体；该基因编辑载体的筛选标记是潮霉素抗性。
7.一种单倍体介导培育Oslg1突变体表型改良水稻株型的方法，其特征在于，包括：(1)将权利要求2所述的编辑OsLG1基因的基因编辑载体以及权利要求5所述的编辑OsMTL基因的基因编辑载体共转化水稻；(2)在共转化的后代中分离挑选出两个OsMTL基因被编辑突变、携带有编辑OsLG1基因的基因编辑载体、但不携带编辑OsMTL基因的基因编辑载体的T2株系；(3)用这两个T2株系的花粉与需要进行基因编辑的水稻材料进行杂交诱导单倍体，获得OsLG1基因发生编辑突变的水稻单倍体植株；(4)将该水稻单倍体进行染色体加倍得到二倍体水稻。
8.按照权利要求7所述的方法，其特征在于：步骤(2)中所述的两个T2株系的筛选方法包括：(a)将共转化后的T0植株用潮霉素和草铵膦除草剂进行筛选，挑选出双阳性的转基因水稻进行自交得到T1代幼苗；(b)用潮霉素和草铵膦除草剂对T1代幼苗进行筛选得到具有草铵膦抗性、但不具有潮霉素抗性的单株；(c)对该单株进行OsMTL基因的测序，找到OsMTL基因发生纯合突变的单株；(d)对这些单株进行自交得到T2代株系。
9.按照权利要求7所述的方法，其特征在于，步骤(2)中所述的两个OsMTL基因被编辑突变后的核苷酸序列分别为SEQ ID No.8和SEQ ID No.9所示；步骤(3)中所述的OsLG1基因被编辑突变后的核苷酸序列为SEQ ID No.10所示。
10.权利要求1或4所述的靶标DNA序列、权利要求2或5所述的基因编辑载体在培育高产水稻新品种或耐密植水稻新品种中的应用。
Priority Applications (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN202010996007.9A CN112522259A (zh) | 2020-09-21 | 2020-09-21 | 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 |
Applications Claiming Priority (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN202010996007.9A CN112522259A (zh) | 2020-09-21 | 2020-09-21 | 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 |
Publications (1)
| Publication Number | Publication Date |
|---|---|
| CN112522259A true CN112522259A (zh) | 2021-03-19 |
Family
ID=74980230
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| CN202010996007.9A Pending CN112522259A (zh) | 2020-09-21 | 2020-09-21 | 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 |
Country Status (1)
| Country | Link |
|---|---|
| CN (1) | CN112522259A (zh) |
Cited By (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN114410650A (zh) * | 2022-01-04 | 2022-04-29 | 广东省农业科学院农业质量标准与监测技术研究所 | 一种水稻盐敏感突变基因SS2、突变体ss2以及应用 |
Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO2018015956A1 (en) * | 2016-07-21 | 2018-01-25 | Kaiima Bio Agritech Ltd. | Compositions and methods for generating a haploid of a target plant |
| CN109943585A (zh) * | 2018-04-12 | 2019-06-28 | 中国水稻研究所 | 一种利用植物杂种优势的方法 |
-
2020
- 2020-09-21 CN CN202010996007.9A patent/CN112522259A/zh active Pending
Patent Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO2018015956A1 (en) * | 2016-07-21 | 2018-01-25 | Kaiima Bio Agritech Ltd. | Compositions and methods for generating a haploid of a target plant |
| CN109943585A (zh) * | 2018-04-12 | 2019-06-28 | 中国水稻研究所 | 一种利用植物杂种优势的方法 |
Non-Patent Citations (5)
| Title |
|---|
| CUILAN SHI ET AL.: "Ubiquitin Specific Protease 15 Has an Important Role in Regulating Grain Width and Size in Rice", 《PLANT PHYSIOL.》 * |
| YUFENG HUA等: "A simple and efficient method for CRISPR/Cas9-induced mutant screening", 《JOURNAL OF GENETICS AND GENOMICS》 * |
| ZUOFENG ZHU ET AL.: "Genetic control of inflorescence architecture during rice domestication", 《NATURE COMMUNICATIONS》 * |
| 方肇勤 等: "《分子生物学技术在中医药研究中的应用》", 30 April 2018, 上海科学技术出版社 * |
| 陈海强等: "植物单倍体诱导技术发展与创新", 《遗传》 * |
Cited By (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN114410650A (zh) * | 2022-01-04 | 2022-04-29 | 广东省农业科学院农业质量标准与监测技术研究所 | 一种水稻盐敏感突变基因SS2、突变体ss2以及应用 |
Similar Documents
| Publication | Publication Date | Title |
|---|---|---|
| CN105602952B (zh) | 一种育性基因及其应用 | |
| CN107267527B (zh) | 雄性育性的保持方法及其应用 | |
| US20200140874A1 (en) | Genome Editing-Based Crop Engineering and Production of Brachytic Plants | |
| US20220396805A1 (en) | Dna sequence for regulating maize leaf angle, and mutant, molecular markers, detection primers, and use thereof | |
| CN110218810B (zh) | 调控玉米雄穗构型的启动子、分子标记及其应用 | |
| KR20210039306A (ko) | CRISPR/Cas9과 gRNA 개별 발현 형질전환 식물체를 이용한 유전자 편집 방법 | |
| CN115605500A (zh) | 控制分生组织大小以改良作物的方法 | |
| CN113832179B (zh) | ZmELF3.1蛋白及其功能缺失突变体在调控作物雄穗分支数中的应用 | |
| CN110713994B (zh) | 一种植物耐逆性相关蛋白TaMAPK3及其编码基因和应用 | |
| CN105695477A (zh) | 雄性不育突变体oss125及其应用 | |
| WO2019129145A1 (en) | Flowering time-regulating gene cmp1 and related constructs and applications thereof | |
| CN112522259A (zh) | 单倍体介导培育具有Oslg1突变体表型的株型改良水稻材料的方法 | |
| CN102477091B (zh) | 水稻雄性不育蛋白及其编码基因与应用 | |
| CN115466747B (zh) | 糖基转移酶ZmKOB1基因及其在调控玉米雌穗结实性状或发育上的应用 | |
| CN113817033B (zh) | ZmELF3.1蛋白及其功能缺失突变体在调控作物气生根数目或层数的应用 | |
| JPH07503850A (ja) | 組み替えジベレリンｄｎａおよびその用途 | |
| CN108660139A (zh) | 植物育性调控基因np2及其编码蛋白和应用 | |
| CN110079535B (zh) | 玉米ZmPIF3s突变型蛋白、其编码基因及其在育种上的应用 | |
| CN113151295A (zh) | 水稻温敏雄性不育基因OsFMS1及其应用 | |
| US7176350B2 (en) | Method for controlling ratio of proteins/lipids in crop seeds | |
| CN114644692B (zh) | 一种定点突变创制旱敏感玉米种质的方法及其应用 | |
| CN113462661B (zh) | 从玉米中分离的siz1蛋白及其编码基因和在品种改良中的应用 | |
| CN114835816B (zh) | 一种调控植物基因组dna特定区域甲基化水平的方法 | |
| CN116789785B (zh) | 长雄蕊野生稻高产、高光效基因FarL1及其应用 | |
| CN113416747B (zh) | 一种创建温度敏感型雄性不育植物的方法 |
Legal Events
| Date | Code | Title | Description |
|---|---|---|---|
| PB01 | Publication | ||
| PB01 | Publication | ||
| SE01 | Entry into force of request for substantive examination | ||
| SE01 | Entry into force of request for substantive examination | ||
| RJ01 | Rejection of invention patent application after publication | ||
| RJ01 | Rejection of invention patent application after publication |
Application publication date: 20210319 |