CN112176083B - 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 - Google Patents
小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 Download PDFInfo
- Publication number
- CN112176083B CN112176083B CN201910602364.XA CN201910602364A CN112176083B CN 112176083 B CN112176083 B CN 112176083B CN 201910602364 A CN201910602364 A CN 201910602364A CN 112176083 B CN112176083 B CN 112176083B
- Authority
- CN
- China
- Prior art keywords
- wheat
- powdery mildew
- sequence
- detected
- dna fragment
- Prior art date
- Legal status (The legal status is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the status listed.)
- Active
Links
- 241000209140 Triticum Species 0.000 title claims abstract description 153
- 235000021307 Triticum Nutrition 0.000 title claims abstract description 153
- 241000221785 Erysiphales Species 0.000 title claims abstract description 136
- 108090000623 proteins and genes Proteins 0.000 title abstract description 36
- 239000003147 molecular marker Substances 0.000 title abstract description 29
- 108020004414 DNA Proteins 0.000 claims abstract description 87
- 239000012634 fragment Substances 0.000 claims abstract description 52
- 102000053602 DNA Human genes 0.000 claims abstract description 7
- 108020004682 Single-Stranded DNA Proteins 0.000 claims abstract description 6
- 238000012408 PCR amplification Methods 0.000 claims description 56
- 238000000034 method Methods 0.000 claims description 46
- 201000010099 disease Diseases 0.000 claims description 42
- 208000037265 diseases, disorders, signs and symptoms Diseases 0.000 claims description 42
- 238000001976 enzyme digestion Methods 0.000 claims description 20
- 108091008146 restriction endonucleases Proteins 0.000 claims description 18
- 238000012216 screening Methods 0.000 claims description 17
- 239000002773 nucleotide Substances 0.000 claims description 15
- 125000003729 nucleotide group Chemical group 0.000 claims description 15
- 238000012163 sequencing technique Methods 0.000 claims description 10
- 239000000463 material Substances 0.000 abstract description 9
- 238000009395 breeding Methods 0.000 abstract description 6
- 230000001488 breeding effect Effects 0.000 abstract description 6
- 101150013841 CNL gene Proteins 0.000 description 41
- 208000015181 infectious disease Diseases 0.000 description 31
- 230000002068 genetic effect Effects 0.000 description 18
- 239000013612 plasmid Substances 0.000 description 17
- 208000035240 Disease Resistance Diseases 0.000 description 15
- 240000008056 Triticum dicoccoides Species 0.000 description 12
- 235000018680 Triticum dicoccoides Nutrition 0.000 description 12
- 238000001514 detection method Methods 0.000 description 10
- 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 9
- 244000098338 Triticum aestivum Species 0.000 description 7
- 239000003153 chemical reaction reagent Substances 0.000 description 7
- 238000010276 construction Methods 0.000 description 7
- 230000009261 transgenic effect Effects 0.000 description 7
- 238000011161 development Methods 0.000 description 6
- 241000196324 Embryophyta Species 0.000 description 5
- 230000000295 complement effect Effects 0.000 description 5
- 230000014509 gene expression Effects 0.000 description 5
- 229920002477 rna polymer Polymers 0.000 description 5
- 241000589158 Agrobacterium Species 0.000 description 4
- 238000006243 chemical reaction Methods 0.000 description 4
- 238000010367 cloning Methods 0.000 description 4
- 239000002299 complementary DNA Substances 0.000 description 4
- 230000006801 homologous recombination Effects 0.000 description 4
- 238000002744 homologous recombination Methods 0.000 description 4
- 238000009396 hybridization Methods 0.000 description 4
- 238000011081 inoculation Methods 0.000 description 4
- 239000003550 marker Substances 0.000 description 4
- 230000009467 reduction Effects 0.000 description 4
- 238000011426 transformation method Methods 0.000 description 4
- 108010085238 Actins Proteins 0.000 description 3
- 241000589155 Agrobacterium tumefaciens Species 0.000 description 3
- 101000855256 Homo sapiens Uncharacterized protein C16orf74 Proteins 0.000 description 3
- 108700019146 Transgenes Proteins 0.000 description 3
- 102100026591 Uncharacterized protein C16orf74 Human genes 0.000 description 3
- 238000004458 analytical method Methods 0.000 description 3
- 238000002474 experimental method Methods 0.000 description 3
- 238000000605 extraction Methods 0.000 description 3
- 238000004519 manufacturing process Methods 0.000 description 3
- 238000013507 mapping Methods 0.000 description 3
- HOVPGJUNRLMIOZ-CIUDSAMLSA-N Ala-Ser-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@H](C)N HOVPGJUNRLMIOZ-CIUDSAMLSA-N 0.000 description 2
- GXXWTNKNFFKTJB-NAKRPEOUSA-N Arg-Ile-Ser Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CO)C(O)=O GXXWTNKNFFKTJB-NAKRPEOUSA-N 0.000 description 2
- 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 2
- RYQSYXFGFOTJDJ-RHYQMDGZSA-N Arg-Thr-Leu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(O)=O RYQSYXFGFOTJDJ-RHYQMDGZSA-N 0.000 description 2
- 241000894006 Bacteria Species 0.000 description 2
- 108091026890 Coding region Proteins 0.000 description 2
- PVBBEKPHARMPHX-DCAQKATOSA-N Glu-Gln-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CCC(N)=O)NC(=O)[C@@H](N)CCC(O)=O PVBBEKPHARMPHX-DCAQKATOSA-N 0.000 description 2
- YSDLIYZLOTZZNP-UWVGGRQHSA-N Gly-Leu-Met Chemical compound CSCC[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)CN YSDLIYZLOTZZNP-UWVGGRQHSA-N 0.000 description 2
- SJLKKOZFHSJJAW-YUMQZZPRSA-N Gly-Met-Glu Chemical compound CSCC[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)O)NC(=O)CN SJLKKOZFHSJJAW-YUMQZZPRSA-N 0.000 description 2
- JRHFQUPIZOYKQP-KBIXCLLPSA-N Ile-Ala-Glu Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCC(O)=O JRHFQUPIZOYKQP-KBIXCLLPSA-N 0.000 description 2
- 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 2
- 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 2
- QVFGXCVIXXBFHO-AVGNSLFASA-N Leu-Glu-Leu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O QVFGXCVIXXBFHO-AVGNSLFASA-N 0.000 description 2
- HGFGEMSVBMCFKK-MNXVOIDGSA-N Leu-Ile-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(O)=O)C(O)=O HGFGEMSVBMCFKK-MNXVOIDGSA-N 0.000 description 2
- WMIOEVKKYIMVKI-DCAQKATOSA-N Leu-Pro-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N1CCC[C@H]1C(=O)N[C@@H](C)C(O)=O WMIOEVKKYIMVKI-DCAQKATOSA-N 0.000 description 2
- ONPDTSFZAIWMDI-AVGNSLFASA-N Lys-Leu-Gln Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O ONPDTSFZAIWMDI-AVGNSLFASA-N 0.000 description 2
- SITLTJHOQZFJGG-UHFFFAOYSA-N N-L-alpha-glutamyl-L-valine Natural products CC(C)C(C(O)=O)NC(=O)C(N)CCC(O)=O SITLTJHOQZFJGG-UHFFFAOYSA-N 0.000 description 2
- DOSZISJPMCYEHT-NAKRPEOUSA-N Ser-Ile-Val Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C(C)C)C(O)=O DOSZISJPMCYEHT-NAKRPEOUSA-N 0.000 description 2
- 108091081024 Start codon Proteins 0.000 description 2
- 238000000246 agarose gel electrophoresis Methods 0.000 description 2
- 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 2
- 230000003321 amplification Effects 0.000 description 2
- 239000007864 aqueous solution Substances 0.000 description 2
- 108010013835 arginine glutamate Proteins 0.000 description 2
- 235000013339 cereals Nutrition 0.000 description 2
- 210000000349 chromosome Anatomy 0.000 description 2
- 230000000052 comparative effect Effects 0.000 description 2
- NKLPQNGYXWVELD-UHFFFAOYSA-M coomassie brilliant blue Chemical compound [Na+].C1=CC(OCC)=CC=C1NC1=CC=C(C(=C2C=CC(C=C2)=[N+](CC)CC=2C=C(C=CC=2)S([O-])(=O)=O)C=2C=CC(=CC=2)N(CC)CC=2C=C(C=CC=2)S([O-])(=O)=O)C=C1 NKLPQNGYXWVELD-UHFFFAOYSA-M 0.000 description 2
- 239000012154 double-distilled water Substances 0.000 description 2
- 238000010195 expression analysis Methods 0.000 description 2
- 235000013305 food Nutrition 0.000 description 2
- 238000003208 gene overexpression Methods 0.000 description 2
- VPZXBVLAVMBEQI-UHFFFAOYSA-N glycyl-DL-alpha-alanine Natural products OC(=O)C(C)NC(=O)CN VPZXBVLAVMBEQI-UHFFFAOYSA-N 0.000 description 2
- 108010036413 histidylglycine Proteins 0.000 description 2
- 108010025306 histidylleucine Proteins 0.000 description 2
- 108010073472 leucyl-prolyl-proline Proteins 0.000 description 2
- 108010057821 leucylproline Proteins 0.000 description 2
- 230000004807 localization Effects 0.000 description 2
- 230000001404 mediated effect Effects 0.000 description 2
- 108010068488 methionylphenylalanine Proteins 0.000 description 2
- 238000003199 nucleic acid amplification method Methods 0.000 description 2
- 230000002018 overexpression Effects 0.000 description 2
- 108010051242 phenylalanylserine Proteins 0.000 description 2
- 238000003753 real-time PCR Methods 0.000 description 2
- 230000001105 regulatory effect Effects 0.000 description 2
- 102200067253 rs762798018 Human genes 0.000 description 2
- 238000009394 selective breeding Methods 0.000 description 2
- 230000035939 shock Effects 0.000 description 2
- 238000010186 staining Methods 0.000 description 2
- 230000001131 transforming effect Effects 0.000 description 2
- 101150084750 1 gene Proteins 0.000 description 1
- 108020005345 3' Untranslated Regions Proteins 0.000 description 1
- 108020003589 5' Untranslated Regions Proteins 0.000 description 1
- 102000007469 Actins Human genes 0.000 description 1
- 241001522110 Aegilops tauschii Species 0.000 description 1
- ODWSTKXGQGYHSH-FXQIFTODSA-N Ala-Arg-Ala Chemical compound C[C@H](N)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](C)C(O)=O ODWSTKXGQGYHSH-FXQIFTODSA-N 0.000 description 1
- DVWVZSJAYIJZFI-FXQIFTODSA-N Ala-Arg-Asn Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(N)=O)C(O)=O DVWVZSJAYIJZFI-FXQIFTODSA-N 0.000 description 1
- TTXMOJWKNRJWQJ-FXQIFTODSA-N Ala-Arg-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)C)CCCN=C(N)N TTXMOJWKNRJWQJ-FXQIFTODSA-N 0.000 description 1
- ZIBWKCRKNFYTPT-ZKWXMUAHSA-N Ala-Asn-Val Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](C(C)C)C(O)=O ZIBWKCRKNFYTPT-ZKWXMUAHSA-N 0.000 description 1
- YIGLXQRFQVWFEY-NRPADANISA-N Ala-Gln-Val Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(O)=O YIGLXQRFQVWFEY-NRPADANISA-N 0.000 description 1
- NBTGEURICRTMGL-WHFBIAKZSA-N Ala-Gly-Ser Chemical compound C[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O NBTGEURICRTMGL-WHFBIAKZSA-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
- 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 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
- MDNAVFBZPROEHO-DCAQKATOSA-N Ala-Lys-Val Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C(C)C)C(O)=O MDNAVFBZPROEHO-DCAQKATOSA-N 0.000 description 1
- MDNAVFBZPROEHO-UHFFFAOYSA-N Ala-Lys-Val Natural products CC(C)C(C(O)=O)NC(=O)C(NC(=O)C(C)N)CCCCN MDNAVFBZPROEHO-UHFFFAOYSA-N 0.000 description 1
- OSRZOHXQCUFIQG-FPMFFAJLSA-N Ala-Phe-Pro Chemical compound C([C@H](NC(=O)[C@@H]([NH3+])C)C(=O)N1[C@H](CCC1)C([O-])=O)C1=CC=CC=C1 OSRZOHXQCUFIQG-FPMFFAJLSA-N 0.000 description 1
- 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 1
- WQKAQKZRDIZYNV-VZFHVOOUSA-N Ala-Ser-Thr Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(O)=O WQKAQKZRDIZYNV-VZFHVOOUSA-N 0.000 description 1
- SYIFFFHSXBNPMC-UWJYBYFXSA-N Ala-Ser-Tyr Chemical compound C[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)O)N SYIFFFHSXBNPMC-UWJYBYFXSA-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
- 101000956243 Arabidopsis thaliana CSC1-like protein HYP1 Proteins 0.000 description 1
- SBVJJNJLFWSJOV-UBHSHLNASA-N Arg-Ala-Phe Chemical compound NC(=N)NCCC[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 SBVJJNJLFWSJOV-UBHSHLNASA-N 0.000 description 1
- UXJCMQFPDWCHKX-DCAQKATOSA-N Arg-Arg-Glu Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCC(O)=O)C(O)=O UXJCMQFPDWCHKX-DCAQKATOSA-N 0.000 description 1
- JTKLCCFLSLCCST-SZMVWBNQSA-N Arg-Arg-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@H](CCCN=C(N)N)N)C(O)=O)=CNC2=C1 JTKLCCFLSLCCST-SZMVWBNQSA-N 0.000 description 1
- OZNSCVPYWZRQPY-CIUDSAMLSA-N Arg-Asp-Glu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O OZNSCVPYWZRQPY-CIUDSAMLSA-N 0.000 description 1
- WVNFNPGXYADPPO-BQBZGAKWSA-N Arg-Gly-Ser Chemical compound NC(N)=NCCC[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O WVNFNPGXYADPPO-BQBZGAKWSA-N 0.000 description 1
- BNYNOWJESJJIOI-XUXIUFHCSA-N Arg-Lys-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](CCCN=C(N)N)N BNYNOWJESJJIOI-XUXIUFHCSA-N 0.000 description 1
- KMFPQTITXUKJOV-DCAQKATOSA-N Arg-Ser-Leu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O KMFPQTITXUKJOV-DCAQKATOSA-N 0.000 description 1
- UVTGNSWSRSCPLP-UHFFFAOYSA-N Arg-Tyr Natural products NC(CCNC(=N)N)C(=O)NC(Cc1ccc(O)cc1)C(=O)O UVTGNSWSRSCPLP-UHFFFAOYSA-N 0.000 description 1
- KSBHCUSPLWRVEK-ZLUOBGJFSA-N Asn-Asn-Asp Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O KSBHCUSPLWRVEK-ZLUOBGJFSA-N 0.000 description 1
- NNMUHYLAYUSTTN-FXQIFTODSA-N Asn-Gln-Glu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O NNMUHYLAYUSTTN-FXQIFTODSA-N 0.000 description 1
- JREOBWLIZLXRIS-GUBZILKMSA-N Asn-Glu-Leu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O JREOBWLIZLXRIS-GUBZILKMSA-N 0.000 description 1
- LVHMEJJWEXBMKK-GMOBBJLQSA-N Asn-Ile-Met Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)O)NC(=O)[C@H](CC(=O)N)N LVHMEJJWEXBMKK-GMOBBJLQSA-N 0.000 description 1
- JLNFZLNDHONLND-GARJFASQSA-N Asn-Leu-Pro Chemical compound CC(C)C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CC(=O)N)N JLNFZLNDHONLND-GARJFASQSA-N 0.000 description 1
- DJIMLSXHXKWADV-CIUDSAMLSA-N Asn-Leu-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H](N)CC(N)=O DJIMLSXHXKWADV-CIUDSAMLSA-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
- BKFXFUPYETWGGA-XVSYOHENSA-N Asn-Phe-Thr Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H]([C@@H](C)O)C(O)=O BKFXFUPYETWGGA-XVSYOHENSA-N 0.000 description 1
- YUOXLJYVSZYPBJ-CIUDSAMLSA-N Asn-Pro-Glu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCC(O)=O)C(O)=O YUOXLJYVSZYPBJ-CIUDSAMLSA-N 0.000 description 1
- VHQSGALUSWIYOD-QXEWZRGKSA-N Asn-Pro-Val Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(O)=O VHQSGALUSWIYOD-QXEWZRGKSA-N 0.000 description 1
- MYTHOBCLNIOFBL-SRVKXCTJSA-N Asn-Ser-Tyr Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O MYTHOBCLNIOFBL-SRVKXCTJSA-N 0.000 description 1
- LMIWYCWRJVMAIQ-NHCYSSNCSA-N Asn-Val-Lys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CC(=O)N)N LMIWYCWRJVMAIQ-NHCYSSNCSA-N 0.000 description 1
- QRULNKJGYQQZMW-ZLUOBGJFSA-N Asp-Asn-Asp Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O QRULNKJGYQQZMW-ZLUOBGJFSA-N 0.000 description 1
- JGDBHIVECJGXJA-FXQIFTODSA-N Asp-Asp-Arg Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O JGDBHIVECJGXJA-FXQIFTODSA-N 0.000 description 1
- QXHVOUSPVAWEMX-ZLUOBGJFSA-N Asp-Asp-Ser Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(O)=O QXHVOUSPVAWEMX-ZLUOBGJFSA-N 0.000 description 1
- WJHYGGVCWREQMO-GHCJXIJMSA-N Asp-Cys-Ile Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CS)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O WJHYGGVCWREQMO-GHCJXIJMSA-N 0.000 description 1
- HRGGPWBIMIQANI-GUBZILKMSA-N Asp-Gln-Leu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O HRGGPWBIMIQANI-GUBZILKMSA-N 0.000 description 1
- RRKCPMGSRIDLNC-AVGNSLFASA-N Asp-Glu-Tyr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O RRKCPMGSRIDLNC-AVGNSLFASA-N 0.000 description 1
- YNCHFVRXEQFPBY-BQBZGAKWSA-N Asp-Gly-Arg Chemical compound OC(=O)C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCN=C(N)N YNCHFVRXEQFPBY-BQBZGAKWSA-N 0.000 description 1
- BIVYLQMZPHDUIH-WHFBIAKZSA-N Asp-Gly-Cys Chemical compound C([C@@H](C(=O)NCC(=O)N[C@@H](CS)C(=O)O)N)C(=O)O BIVYLQMZPHDUIH-WHFBIAKZSA-N 0.000 description 1
- PZXPWHFYZXTFBI-YUMQZZPRSA-N Asp-Gly-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CC(O)=O PZXPWHFYZXTFBI-YUMQZZPRSA-N 0.000 description 1
- TVIZQBFURPLQDV-DJFWLOJKSA-N Asp-His-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CC1=CN=CN1)NC(=O)[C@H](CC(=O)O)N TVIZQBFURPLQDV-DJFWLOJKSA-N 0.000 description 1
- ICZWAZVKLACMKR-CIUDSAMLSA-N Asp-His-Ser Chemical compound OC(=O)C[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](CO)C(O)=O)CC1=CN=CN1 ICZWAZVKLACMKR-CIUDSAMLSA-N 0.000 description 1
- UZNSWMFLKVKJLI-VHWLVUOQSA-N Asp-Ile-Trp Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(O)=O UZNSWMFLKVKJLI-VHWLVUOQSA-N 0.000 description 1
- CJUKAWUWBZCTDQ-SRVKXCTJSA-N Asp-Leu-Lys Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(O)=O CJUKAWUWBZCTDQ-SRVKXCTJSA-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
- UZFHNLYQWMGUHU-DCAQKATOSA-N Asp-Lys-Arg Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O UZFHNLYQWMGUHU-DCAQKATOSA-N 0.000 description 1
- RXBGWGRSWXOBGK-KKUMJFAQSA-N Asp-Lys-Tyr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O RXBGWGRSWXOBGK-KKUMJFAQSA-N 0.000 description 1
- VMVUDJUXJKDGNR-FXQIFTODSA-N Asp-Met-Asn Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CC(=O)O)N VMVUDJUXJKDGNR-FXQIFTODSA-N 0.000 description 1
- YRZIYQGXTSBRLT-AVGNSLFASA-N Asp-Phe-Gln Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O YRZIYQGXTSBRLT-AVGNSLFASA-N 0.000 description 1
- ZKAOJVJQGVUIIU-GUBZILKMSA-N Asp-Pro-Arg Chemical compound OC(=O)C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCCNC(N)=N)C(O)=O ZKAOJVJQGVUIIU-GUBZILKMSA-N 0.000 description 1
- BWJZSLQJNBSUPM-FXQIFTODSA-N Asp-Pro-Asn Chemical compound OC(=O)C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC(N)=O)C(O)=O BWJZSLQJNBSUPM-FXQIFTODSA-N 0.000 description 1
- ZBYLEBZCVKLPCY-FXQIFTODSA-N Asp-Ser-Arg Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O ZBYLEBZCVKLPCY-FXQIFTODSA-N 0.000 description 1
- NBKLEMWHDLAUEM-CIUDSAMLSA-N Asp-Ser-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CC(=O)O)N NBKLEMWHDLAUEM-CIUDSAMLSA-N 0.000 description 1
- QSFHZPQUAAQHAQ-CIUDSAMLSA-N Asp-Ser-Leu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O QSFHZPQUAAQHAQ-CIUDSAMLSA-N 0.000 description 1
- DZSICRGTVPDCRN-YUMQZZPRSA-N Cys-Gly-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CS)N DZSICRGTVPDCRN-YUMQZZPRSA-N 0.000 description 1
- WVLZTXGTNGHPBO-SRVKXCTJSA-N Cys-Leu-Leu Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O WVLZTXGTNGHPBO-SRVKXCTJSA-N 0.000 description 1
- CIVXDCMSSFGWAL-YUMQZZPRSA-N Cys-Lys-Gly Chemical compound C(CCN)C[C@@H](C(=O)NCC(=O)O)NC(=O)[C@H](CS)N CIVXDCMSSFGWAL-YUMQZZPRSA-N 0.000 description 1
- ZXCAQANTQWBICD-DCAQKATOSA-N Cys-Lys-Val Chemical compound CC(C)[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](CS)N ZXCAQANTQWBICD-DCAQKATOSA-N 0.000 description 1
- KJJASVYBTKRYSN-FXQIFTODSA-N Cys-Pro-Asp Chemical compound C1C[C@H](N(C1)C(=O)[C@H](CS)N)C(=O)N[C@@H](CC(=O)O)C(=O)O KJJASVYBTKRYSN-FXQIFTODSA-N 0.000 description 1
- UEHCDNYDBBCQEL-CIUDSAMLSA-N Cys-Ser-His Chemical compound C1=C(NC=N1)C[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CS)N UEHCDNYDBBCQEL-CIUDSAMLSA-N 0.000 description 1
- JLZCAZJGWNRXCI-XKBZYTNZSA-N Cys-Thr-Glu Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(O)=O JLZCAZJGWNRXCI-XKBZYTNZSA-N 0.000 description 1
- ZFHXNNXMNLWKJH-HJPIBITLSA-N Cys-Tyr-Ile Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O ZFHXNNXMNLWKJH-HJPIBITLSA-N 0.000 description 1
- 108010090461 DFG peptide Proteins 0.000 description 1
- 108700024394 Exon Proteins 0.000 description 1
- 241000233866 Fungi Species 0.000 description 1
- 108700039691 Genetic Promoter Regions Proteins 0.000 description 1
- LTLXPHKSQQILNF-CIUDSAMLSA-N Gln-Arg-Cys Chemical compound C(C[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CCC(=O)N)N)CN=C(N)N LTLXPHKSQQILNF-CIUDSAMLSA-N 0.000 description 1
- 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 1
- MFJAPSYJQJCQDN-BQBZGAKWSA-N Gln-Gly-Glu Chemical compound NC(=O)CC[C@H](N)C(=O)NCC(=O)N[C@@H](CCC(O)=O)C(O)=O MFJAPSYJQJCQDN-BQBZGAKWSA-N 0.000 description 1
- UXXIVIQGOODKQC-NUMRIWBASA-N Gln-Thr-Asn Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CCC(=O)N)N)O UXXIVIQGOODKQC-NUMRIWBASA-N 0.000 description 1
- OACPJRQRAHMQEQ-NHCYSSNCSA-N Gln-Val-Arg Chemical compound NC(=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 OACPJRQRAHMQEQ-NHCYSSNCSA-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
- INGJLBQKTRJLFO-UKJIMTQDSA-N Glu-Ile-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H]([C@@H](C)CC)NC(=O)[C@@H](N)CCC(O)=O INGJLBQKTRJLFO-UKJIMTQDSA-N 0.000 description 1
- IRXNJYPKBVERCW-DCAQKATOSA-N Glu-Leu-Glu Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O IRXNJYPKBVERCW-DCAQKATOSA-N 0.000 description 1
- VGBSZQSKQRMLHD-MNXVOIDGSA-N Glu-Leu-Ile Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O VGBSZQSKQRMLHD-MNXVOIDGSA-N 0.000 description 1
- BCYGDJXHAGZNPQ-DCAQKATOSA-N Glu-Lys-Glu Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O BCYGDJXHAGZNPQ-DCAQKATOSA-N 0.000 description 1
- MFNUFCFRAZPJFW-JYJNAYRXSA-N Glu-Lys-Phe Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 MFNUFCFRAZPJFW-JYJNAYRXSA-N 0.000 description 1
- RBXSZQRSEGYDFG-GUBZILKMSA-N Glu-Lys-Ser Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(O)=O RBXSZQRSEGYDFG-GUBZILKMSA-N 0.000 description 1
- ZQYZDDXTNQXUJH-CIUDSAMLSA-N Glu-Met-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCSC)NC(=O)[C@H](CCC(=O)O)N ZQYZDDXTNQXUJH-CIUDSAMLSA-N 0.000 description 1
- LKOAAMXDJGEYMS-ZPFDUUQYSA-N Glu-Met-Ile Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCSC)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O LKOAAMXDJGEYMS-ZPFDUUQYSA-N 0.000 description 1
- CBEUFCJRFNZMCU-SRVKXCTJSA-N Glu-Met-Leu Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O CBEUFCJRFNZMCU-SRVKXCTJSA-N 0.000 description 1
- CHDWDBPJOZVZSE-KKUMJFAQSA-N Glu-Phe-Met Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCSC)C(O)=O CHDWDBPJOZVZSE-KKUMJFAQSA-N 0.000 description 1
- CBOVGULVQSVMPT-CIUDSAMLSA-N Glu-Pro-Asn Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC(N)=O)C(O)=O CBOVGULVQSVMPT-CIUDSAMLSA-N 0.000 description 1
- 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 1
- BDISFWMLMNBTGP-NUMRIWBASA-N Glu-Thr-Asp Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(O)=O BDISFWMLMNBTGP-NUMRIWBASA-N 0.000 description 1
- PHONXOACARQMPM-BQBZGAKWSA-N Gly-Ala-Met Chemical compound [H]NCC(=O)N[C@@H](C)C(=O)N[C@@H](CCSC)C(O)=O PHONXOACARQMPM-BQBZGAKWSA-N 0.000 description 1
- XCLCVBYNGXEVDU-WHFBIAKZSA-N Gly-Asn-Ser Chemical compound NCC(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(O)=O XCLCVBYNGXEVDU-WHFBIAKZSA-N 0.000 description 1
- JUGQPPOVWXSPKJ-RYUDHWBXSA-N Gly-Gln-Phe Chemical compound [H]NCC(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O JUGQPPOVWXSPKJ-RYUDHWBXSA-N 0.000 description 1
- HDNXXTBKOJKWNN-WDSKDSINSA-N Gly-Glu-Asn Chemical compound NCC(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O HDNXXTBKOJKWNN-WDSKDSINSA-N 0.000 description 1
- YYPFZVIXAVDHIK-IUCAKERBSA-N Gly-Glu-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CCC(O)=O)NC(=O)CN YYPFZVIXAVDHIK-IUCAKERBSA-N 0.000 description 1
- UTYGDAHJBBDPBA-BYULHYEWSA-N Gly-Ile-Asp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)CN UTYGDAHJBBDPBA-BYULHYEWSA-N 0.000 description 1
- HAXARWKYFIIHKD-ZKWXMUAHSA-N Gly-Ile-Ser Chemical compound NCC(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CO)C(O)=O HAXARWKYFIIHKD-ZKWXMUAHSA-N 0.000 description 1
- CCBIBMKQNXHNIN-ZETCQYMHSA-N Gly-Leu-Gly Chemical compound NCC(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O CCBIBMKQNXHNIN-ZETCQYMHSA-N 0.000 description 1
- UUYBFNKHOCJCHT-VHSXEESVSA-N Gly-Leu-Pro Chemical compound CC(C)C[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)CN UUYBFNKHOCJCHT-VHSXEESVSA-N 0.000 description 1
- IBYOLNARKHMLBG-WHOFXGATSA-N Gly-Phe-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)CN)CC1=CC=CC=C1 IBYOLNARKHMLBG-WHOFXGATSA-N 0.000 description 1
- CSMYMGFCEJWALV-WDSKDSINSA-N Gly-Ser-Gln Chemical compound NCC(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCC(N)=O CSMYMGFCEJWALV-WDSKDSINSA-N 0.000 description 1
- DBUNZBWUWCIELX-JHEQGTHGSA-N Gly-Thr-Glu Chemical compound [H]NCC(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(O)=O DBUNZBWUWCIELX-JHEQGTHGSA-N 0.000 description 1
- HQSKKSLNLSTONK-JTQLQIEISA-N Gly-Tyr-Gly Chemical compound OC(=O)CNC(=O)[C@@H](NC(=O)CN)CC1=CC=C(O)C=C1 HQSKKSLNLSTONK-JTQLQIEISA-N 0.000 description 1
- 102000051366 Glycosyltransferases Human genes 0.000 description 1
- 108700023372 Glycosyltransferases Proteins 0.000 description 1
- 241000896246 Golovinomyces cichoracearum Species 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
- 241001000287 Helvetia Species 0.000 description 1
- PDSUIXMZYNURGI-AVGNSLFASA-N His-Arg-Arg Chemical compound NC(N)=NCCC[C@@H](C(O)=O)NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@@H](N)CC1=CN=CN1 PDSUIXMZYNURGI-AVGNSLFASA-N 0.000 description 1
- WJGSTIMGSIWHJX-HVTMNAMFSA-N His-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](CC1=CN=CN1)N WJGSTIMGSIWHJX-HVTMNAMFSA-N 0.000 description 1
- OQDLKDUVMTUPPG-AVGNSLFASA-N His-Leu-Glu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O OQDLKDUVMTUPPG-AVGNSLFASA-N 0.000 description 1
- BPOHQCZZSFBSON-KKUMJFAQSA-N His-Leu-His Chemical compound CC(C)C[C@H](NC(=O)[C@@H](N)Cc1cnc[nH]1)C(=O)N[C@@H](Cc1cnc[nH]1)C(O)=O BPOHQCZZSFBSON-KKUMJFAQSA-N 0.000 description 1
- GJMHMDKCJPQJOI-IHRRRGAJSA-N His-Lys-Arg Chemical compound NC(N)=NCCC[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CC1=CN=CN1 GJMHMDKCJPQJOI-IHRRRGAJSA-N 0.000 description 1
- UXSATKFPUVZVDK-KKUMJFAQSA-N His-Lys-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](CC1=CN=CN1)N UXSATKFPUVZVDK-KKUMJFAQSA-N 0.000 description 1
- YOTNPRLPIPHQSB-XUXIUFHCSA-N Ile-Arg-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCCCN)C(=O)O)N YOTNPRLPIPHQSB-XUXIUFHCSA-N 0.000 description 1
- YKRIXHPEIZUDDY-GMOBBJLQSA-N Ile-Asn-Arg Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@H](C(O)=O)CCCN=C(N)N YKRIXHPEIZUDDY-GMOBBJLQSA-N 0.000 description 1
- HDODQNPMSHDXJT-GHCJXIJMSA-N Ile-Asn-Ser Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(O)=O HDODQNPMSHDXJT-GHCJXIJMSA-N 0.000 description 1
- RGSOCXHDOPQREB-ZPFDUUQYSA-N Ile-Asp-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CC(C)C)C(=O)O)N RGSOCXHDOPQREB-ZPFDUUQYSA-N 0.000 description 1
- OVPYIUNCVSOVNF-ZPFDUUQYSA-N Ile-Gln-Pro Natural products CC[C@H](C)[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N1CCC[C@H]1C(O)=O OVPYIUNCVSOVNF-ZPFDUUQYSA-N 0.000 description 1
- JDAWAWXGAUZPNJ-ZPFDUUQYSA-N Ile-Glu-Arg Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)N JDAWAWXGAUZPNJ-ZPFDUUQYSA-N 0.000 description 1
- SPQWWEZBHXHUJN-KBIXCLLPSA-N Ile-Glu-Ser Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O SPQWWEZBHXHUJN-KBIXCLLPSA-N 0.000 description 1
- IGJWJGIHUFQANP-LAEOZQHASA-N Ile-Gly-Gln Chemical compound CC[C@H](C)[C@@H](C(=O)NCC(=O)N[C@@H](CCC(=O)N)C(=O)O)N IGJWJGIHUFQANP-LAEOZQHASA-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
- SJLVSMMIFYTSGY-GRLWGSQLSA-N Ile-Ile-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N SJLVSMMIFYTSGY-GRLWGSQLSA-N 0.000 description 1
- PKGGWLOLRLOPGK-XUXIUFHCSA-N Ile-Leu-Arg Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CCCN=C(N)N PKGGWLOLRLOPGK-XUXIUFHCSA-N 0.000 description 1
- FZWVCYCYWCLQDH-NHCYSSNCSA-N Ile-Leu-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)NCC(=O)O)N FZWVCYCYWCLQDH-NHCYSSNCSA-N 0.000 description 1
- IOVUXUSIGXCREV-DKIMLUQUSA-N Ile-Leu-Phe Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 IOVUXUSIGXCREV-DKIMLUQUSA-N 0.000 description 1
- RFMDODRWJZHZCR-BJDJZHNGSA-N Ile-Lys-Cys Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CS)C(O)=O RFMDODRWJZHZCR-BJDJZHNGSA-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
- PARSHQDZROHERM-NHCYSSNCSA-N Ile-Lys-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)NCC(=O)O)N PARSHQDZROHERM-NHCYSSNCSA-N 0.000 description 1
- FFJQAEYLAQMGDL-MGHWNKPDSA-N Ile-Lys-Tyr Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 FFJQAEYLAQMGDL-MGHWNKPDSA-N 0.000 description 1
- BKPPWVSPSIUXHZ-OSUNSFLBSA-N Ile-Met-Thr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)N[C@@H]([C@@H](C)O)C(=O)O)N BKPPWVSPSIUXHZ-OSUNSFLBSA-N 0.000 description 1
- SNHYFFQZRFIRHO-CYDGBPFRSA-N Ile-Met-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)N[C@@H](C(C)C)C(=O)O)N SNHYFFQZRFIRHO-CYDGBPFRSA-N 0.000 description 1
- IIWQTXMUALXGOV-PCBIJLKTSA-N Ile-Phe-Asp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(=O)O)C(=O)O)N IIWQTXMUALXGOV-PCBIJLKTSA-N 0.000 description 1
- UAELWXJFLZBKQS-WHOFXGATSA-N Ile-Phe-Gly Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](Cc1ccccc1)C(=O)NCC(O)=O UAELWXJFLZBKQS-WHOFXGATSA-N 0.000 description 1
- KBDIBHQICWDGDL-PPCPHDFISA-N Ile-Thr-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(=O)O)N KBDIBHQICWDGDL-PPCPHDFISA-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
- 241000880493 Leptailurus serval Species 0.000 description 1
- CQQGCWPXDHTTNF-GUBZILKMSA-N Leu-Ala-Glu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCC(O)=O CQQGCWPXDHTTNF-GUBZILKMSA-N 0.000 description 1
- QPRQGENIBFLVEB-BJDJZHNGSA-N Leu-Ala-Ile Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](C)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O QPRQGENIBFLVEB-BJDJZHNGSA-N 0.000 description 1
- CUXRXAIAVYLVFD-ULQDDVLXSA-N Leu-Arg-Tyr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 CUXRXAIAVYLVFD-ULQDDVLXSA-N 0.000 description 1
- VIWUBXKCYJGNCL-SRVKXCTJSA-N Leu-Asn-His Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@H](C(O)=O)CC1=CN=CN1 VIWUBXKCYJGNCL-SRVKXCTJSA-N 0.000 description 1
- WXHFZJFZWNCDNB-KKUMJFAQSA-N Leu-Asn-Tyr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 WXHFZJFZWNCDNB-KKUMJFAQSA-N 0.000 description 1
- VPKIQULSKFVCSM-SRVKXCTJSA-N Leu-Gln-Arg Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O VPKIQULSKFVCSM-SRVKXCTJSA-N 0.000 description 1
- ZTLGVASZOIKNIX-DCAQKATOSA-N Leu-Gln-Glu Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N ZTLGVASZOIKNIX-DCAQKATOSA-N 0.000 description 1
- LOLUPZNNADDTAA-AVGNSLFASA-N Leu-Gln-Leu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O LOLUPZNNADDTAA-AVGNSLFASA-N 0.000 description 1
- QDSKNVXKLPQNOJ-GVXVVHGQSA-N Leu-Gln-Val Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(O)=O QDSKNVXKLPQNOJ-GVXVVHGQSA-N 0.000 description 1
- ZFNLIDNJUWNIJL-WDCWCFNPSA-N Leu-Glu-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O ZFNLIDNJUWNIJL-WDCWCFNPSA-N 0.000 description 1
- AUBMZAMQCOYSIC-MNXVOIDGSA-N Leu-Ile-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(N)=O)C(O)=O AUBMZAMQCOYSIC-MNXVOIDGSA-N 0.000 description 1
- UBZGNBKMIJHOHL-BZSNNMDCSA-N Leu-Leu-Phe Chemical compound CC(C)C[C@H]([NH3+])C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C([O-])=O)CC1=CC=CC=C1 UBZGNBKMIJHOHL-BZSNNMDCSA-N 0.000 description 1
- 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 1
- REPBGZHJKYWFMJ-KKUMJFAQSA-N Leu-Lys-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N REPBGZHJKYWFMJ-KKUMJFAQSA-N 0.000 description 1
- 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 1
- VCHVSKNMTXWIIP-SRVKXCTJSA-N Leu-Lys-Ser Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(O)=O VCHVSKNMTXWIIP-SRVKXCTJSA-N 0.000 description 1
- OVZLLFONXILPDZ-VOAKCMCISA-N Leu-Lys-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)O)C(O)=O OVZLLFONXILPDZ-VOAKCMCISA-N 0.000 description 1
- LZHJZLHSRGWBBE-IHRRRGAJSA-N Leu-Lys-Val Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C(C)C)C(O)=O LZHJZLHSRGWBBE-IHRRRGAJSA-N 0.000 description 1
- GNRPTBRHRRZCMA-RWMBFGLXSA-N Leu-Met-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCSC)C(=O)N1CCC[C@@H]1C(=O)O)N GNRPTBRHRRZCMA-RWMBFGLXSA-N 0.000 description 1
- MVVSHHJKJRZVNY-ACRUOGEOSA-N Leu-Phe-Tyr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O MVVSHHJKJRZVNY-ACRUOGEOSA-N 0.000 description 1
- 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 1
- CHJKEDSZNSONPS-DCAQKATOSA-N Leu-Pro-Ser Chemical compound [H]N[C@@H](CC(C)C)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O CHJKEDSZNSONPS-DCAQKATOSA-N 0.000 description 1
- MVHXGBZUJLWZOH-BJDJZHNGSA-N Leu-Ser-Ile Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O MVHXGBZUJLWZOH-BJDJZHNGSA-N 0.000 description 1
- BRTVHXHCUSXYRI-CIUDSAMLSA-N Leu-Ser-Ser Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CO)C(O)=O BRTVHXHCUSXYRI-CIUDSAMLSA-N 0.000 description 1
- PPGBXYKMUMHFBF-KATARQTJSA-N Leu-Ser-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(O)=O PPGBXYKMUMHFBF-KATARQTJSA-N 0.000 description 1
- SQUFDMCWMFOEBA-KKUMJFAQSA-N Leu-Ser-Tyr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 SQUFDMCWMFOEBA-KKUMJFAQSA-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
- XZNJZXJZBMBGGS-NHCYSSNCSA-N Leu-Val-Asn Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(N)=O)C(O)=O XZNJZXJZBMBGGS-NHCYSSNCSA-N 0.000 description 1
- 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 1
- QESXLSQLQHHTIX-RHYQMDGZSA-N Leu-Val-Thr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O QESXLSQLQHHTIX-RHYQMDGZSA-N 0.000 description 1
- NTXYXFDMIHXTHE-WDSOQIARSA-N Leu-Val-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CC(C)C)C(O)=O)=CNC2=C1 NTXYXFDMIHXTHE-WDSOQIARSA-N 0.000 description 1
- HKCCVDWHHTVVPN-CIUDSAMLSA-N Lys-Asp-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C)C(O)=O HKCCVDWHHTVVPN-CIUDSAMLSA-N 0.000 description 1
- DCRWPTBMWMGADO-AVGNSLFASA-N Lys-Glu-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O DCRWPTBMWMGADO-AVGNSLFASA-N 0.000 description 1
- 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 1
- 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 1
- VSTNAUBHKQPVJX-IHRRRGAJSA-N Lys-Met-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O VSTNAUBHKQPVJX-IHRRRGAJSA-N 0.000 description 1
- LNMKRJJLEFASGA-BZSNNMDCSA-N Lys-Phe-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(C)C)C(O)=O LNMKRJJLEFASGA-BZSNNMDCSA-N 0.000 description 1
- JOSAKOKSPXROGQ-BJDJZHNGSA-N Lys-Ser-Ile Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O JOSAKOKSPXROGQ-BJDJZHNGSA-N 0.000 description 1
- GIKFNMZSGYAPEJ-HJGDQZAQSA-N Lys-Thr-Asp Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(O)=O GIKFNMZSGYAPEJ-HJGDQZAQSA-N 0.000 description 1
- CAVRAQIDHUPECU-UVOCVTCTSA-N Lys-Thr-Thr Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O CAVRAQIDHUPECU-UVOCVTCTSA-N 0.000 description 1
- IMDJSVBFQKDDEQ-MGHWNKPDSA-N Lys-Tyr-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CC1=CC=C(C=C1)O)NC(=O)[C@H](CCCCN)N IMDJSVBFQKDDEQ-MGHWNKPDSA-N 0.000 description 1
- DRRXXZBXDMLGFC-IHRRRGAJSA-N Lys-Val-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CCCCN DRRXXZBXDMLGFC-IHRRRGAJSA-N 0.000 description 1
- IKXQOBUBZSOWDY-AVGNSLFASA-N Lys-Val-Val Chemical compound CC(C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)O)NC(=O)[C@H](CCCCN)N IKXQOBUBZSOWDY-AVGNSLFASA-N 0.000 description 1
- CAODKDAPYGUMLK-FXQIFTODSA-N Met-Asn-Ser Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(O)=O CAODKDAPYGUMLK-FXQIFTODSA-N 0.000 description 1
- KQBJYJXPZBNEIK-DCAQKATOSA-N Met-Glu-Arg Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CCCNC(N)=N KQBJYJXPZBNEIK-DCAQKATOSA-N 0.000 description 1
- DYTWOWJWJCBFLE-IHRRRGAJSA-N Met-His-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)CCSC)CC1=CNC=N1 DYTWOWJWJCBFLE-IHRRRGAJSA-N 0.000 description 1
- BEZJTLKUMFMITF-AVGNSLFASA-N Met-Lys-Arg Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@H](C(O)=O)CCCNC(N)=N BEZJTLKUMFMITF-AVGNSLFASA-N 0.000 description 1
- XPVCDCMPKCERFT-GUBZILKMSA-N Met-Ser-Arg Chemical compound [H]N[C@@H](CCSC)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O XPVCDCMPKCERFT-GUBZILKMSA-N 0.000 description 1
- 208000031888 Mycoses Diseases 0.000 description 1
- 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 1
- 101100442582 Neurospora crassa (strain ATCC 24698 / 74-OR23-1A / CBS 708.71 / DSM 1257 / FGSC 987) spe-1 gene Proteins 0.000 description 1
- 108091028043 Nucleic acid sequence Proteins 0.000 description 1
- MQVFHOPCKNTHGT-MELADBBJSA-N Phe-Asp-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC(=O)O)NC(=O)[C@H](CC2=CC=CC=C2)N)C(=O)O MQVFHOPCKNTHGT-MELADBBJSA-N 0.000 description 1
- HGNGAMWHGGANAU-WHOFXGATSA-N Phe-Gly-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CC1=CC=CC=C1 HGNGAMWHGGANAU-WHOFXGATSA-N 0.000 description 1
- APJPXSFJBMMOLW-KBPBESRZSA-N Phe-Gly-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CC1=CC=CC=C1 APJPXSFJBMMOLW-KBPBESRZSA-N 0.000 description 1
- CWFGECHCRMGPPT-MXAVVETBSA-N Phe-Ile-Ser Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CO)C(O)=O CWFGECHCRMGPPT-MXAVVETBSA-N 0.000 description 1
- LRBSWBVUCLLRLU-BZSNNMDCSA-N Phe-Leu-Lys Chemical compound CC(C)C[C@H](NC(=O)[C@@H](N)Cc1ccccc1)C(=O)N[C@@H](CCCCN)C(O)=O LRBSWBVUCLLRLU-BZSNNMDCSA-N 0.000 description 1
- DNAXXTQSTKOHFO-QEJZJMRPSA-N Phe-Lys-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CC1=CC=CC=C1 DNAXXTQSTKOHFO-QEJZJMRPSA-N 0.000 description 1
- CJAHQEZWDZNSJO-KKUMJFAQSA-N Phe-Lys-Cys Chemical compound NCCCC[C@@H](C(=O)N[C@@H](CS)C(O)=O)NC(=O)[C@@H](N)CC1=CC=CC=C1 CJAHQEZWDZNSJO-KKUMJFAQSA-N 0.000 description 1
- GPSMLZQVIIYLDK-ULQDDVLXSA-N Phe-Lys-Val Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C(C)C)C(O)=O GPSMLZQVIIYLDK-ULQDDVLXSA-N 0.000 description 1
- MCIXMYKSPQUMJG-SRVKXCTJSA-N Phe-Ser-Ser Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CO)C(=O)N[C@@H](CO)C(O)=O MCIXMYKSPQUMJG-SRVKXCTJSA-N 0.000 description 1
- FXEKNHAJIMHRFJ-ULQDDVLXSA-N Phe-Val-His Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)NC(=O)[C@H](CC2=CC=CC=C2)N FXEKNHAJIMHRFJ-ULQDDVLXSA-N 0.000 description 1
- 108020005120 Plant DNA Proteins 0.000 description 1
- 108700001094 Plant Genes Proteins 0.000 description 1
- 241000209504 Poaceae Species 0.000 description 1
- OYEUSRAZOGIDBY-JYJNAYRXSA-N Pro-Arg-Tyr Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O OYEUSRAZOGIDBY-JYJNAYRXSA-N 0.000 description 1
- WGAQWMRJUFQXMF-ZPFDUUQYSA-N Pro-Gln-Ile Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O WGAQWMRJUFQXMF-ZPFDUUQYSA-N 0.000 description 1
- KIPIKSXPPLABPN-CIUDSAMLSA-N Pro-Glu-Asn Chemical compound NC(=O)C[C@@H](C(O)=O)NC(=O)[C@H](CCC(O)=O)NC(=O)[C@@H]1CCCN1 KIPIKSXPPLABPN-CIUDSAMLSA-N 0.000 description 1
- STASJMBVVHNWCG-IHRRRGAJSA-N Pro-His-Leu Chemical compound C([C@@H](C(=O)N[C@@H](CC(C)C)C([O-])=O)NC(=O)[C@H]1[NH2+]CCC1)C1=CN=CN1 STASJMBVVHNWCG-IHRRRGAJSA-N 0.000 description 1
- AQGUSRZKDZYGGV-GMOBBJLQSA-N Pro-Ile-Asp Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(O)=O)C(O)=O AQGUSRZKDZYGGV-GMOBBJLQSA-N 0.000 description 1
- ABSSTGUCBCDKMU-UWVGGRQHSA-N Pro-Lys-Gly Chemical compound NCCCC[C@@H](C(=O)NCC(O)=O)NC(=O)[C@@H]1CCCN1 ABSSTGUCBCDKMU-UWVGGRQHSA-N 0.000 description 1
- RFWXYTJSVDUBBZ-DCAQKATOSA-N Pro-Pro-Glu Chemical compound OC(=O)CC[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@H]1NCCC1 RFWXYTJSVDUBBZ-DCAQKATOSA-N 0.000 description 1
- 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 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
- SHTKRJHDMNSKRM-ULQDDVLXSA-N Pro-Tyr-His Chemical compound C1C[C@H](NC1)C(=O)N[C@@H](CC2=CC=C(C=C2)O)C(=O)N[C@@H](CC3=CN=CN3)C(=O)O SHTKRJHDMNSKRM-ULQDDVLXSA-N 0.000 description 1
- 108050003422 Rare lipoprotein A 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
- DWUIECHTAMYEFL-XVYDVKMFSA-N Ser-Ala-His Chemical compound OC[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CC1=CN=CN1 DWUIECHTAMYEFL-XVYDVKMFSA-N 0.000 description 1
- HRNQLKCLPVKZNE-CIUDSAMLSA-N Ser-Ala-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(O)=O HRNQLKCLPVKZNE-CIUDSAMLSA-N 0.000 description 1
- KAAPNMOKUUPKOE-SRVKXCTJSA-N Ser-Asn-Phe Chemical compound OC[C@H](N)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 KAAPNMOKUUPKOE-SRVKXCTJSA-N 0.000 description 1
- SNNSYBWPPVAXQW-ZLUOBGJFSA-N Ser-Cys-Cys Chemical compound C([C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CS)C(=O)O)N)O SNNSYBWPPVAXQW-ZLUOBGJFSA-N 0.000 description 1
- MOVJSUIKUNCVMG-ZLUOBGJFSA-N Ser-Cys-Ser Chemical compound C([C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CO)C(=O)O)N)O MOVJSUIKUNCVMG-ZLUOBGJFSA-N 0.000 description 1
- MAWSJXHRLWVJEZ-ACZMJKKPSA-N Ser-Gln-Cys Chemical compound C(CC(=O)N)[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CO)N MAWSJXHRLWVJEZ-ACZMJKKPSA-N 0.000 description 1
- SMIDBHKWSYUBRZ-ACZMJKKPSA-N Ser-Glu-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O SMIDBHKWSYUBRZ-ACZMJKKPSA-N 0.000 description 1
- PVDTYLHUWAEYGY-CIUDSAMLSA-N Ser-Glu-Arg Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O PVDTYLHUWAEYGY-CIUDSAMLSA-N 0.000 description 1
- SFTZWNJFZYOLBD-ZDLURKLDSA-N Ser-Gly-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CO SFTZWNJFZYOLBD-ZDLURKLDSA-N 0.000 description 1
- DJACUBDEDBZKLQ-KBIXCLLPSA-N Ser-Ile-Glu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(O)=O)C(O)=O DJACUBDEDBZKLQ-KBIXCLLPSA-N 0.000 description 1
- IFPBAGJBHSNYPR-ZKWXMUAHSA-N Ser-Ile-Gly Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)NCC(O)=O IFPBAGJBHSNYPR-ZKWXMUAHSA-N 0.000 description 1
- GJFYFGOEWLDQGW-GUBZILKMSA-N Ser-Leu-Gln Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)O)NC(=O)[C@H](CO)N GJFYFGOEWLDQGW-GUBZILKMSA-N 0.000 description 1
- XNCUYZKGQOCOQH-YUMQZZPRSA-N Ser-Leu-Gly Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O XNCUYZKGQOCOQH-YUMQZZPRSA-N 0.000 description 1
- HEUVHBXOVZONPU-BJDJZHNGSA-N Ser-Leu-Ile Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O HEUVHBXOVZONPU-BJDJZHNGSA-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
- BCAVNDNYOGTQMQ-AAEUAGOBSA-N Ser-Trp-Gly Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)NCC(O)=O BCAVNDNYOGTQMQ-AAEUAGOBSA-N 0.000 description 1
- FRPNVPKQVFHSQY-BPUTZDHNSA-N Ser-Trp-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@H](CC1=CNC2=CC=CC=C21)NC(=O)[C@H](CO)N FRPNVPKQVFHSQY-BPUTZDHNSA-N 0.000 description 1
- ZVBCMFDJIMUELU-BZSNNMDCSA-N Ser-Tyr-Phe Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)O)NC(=O)[C@H](CC2=CC=C(C=C2)O)NC(=O)[C@H](CO)N ZVBCMFDJIMUELU-BZSNNMDCSA-N 0.000 description 1
- OQSQCUWQOIHECT-YJRXYDGGSA-N Ser-Tyr-Thr Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H]([C@@H](C)O)C(O)=O OQSQCUWQOIHECT-YJRXYDGGSA-N 0.000 description 1
- 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 1
- LGIMRDKGABDMBN-DCAQKATOSA-N Ser-Val-Lys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CO)N LGIMRDKGABDMBN-DCAQKATOSA-N 0.000 description 1
- 208000035199 Tetraploidy Diseases 0.000 description 1
- FQPQPTHMHZKGFM-XQXXSGGOSA-N Thr-Ala-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(O)=O FQPQPTHMHZKGFM-XQXXSGGOSA-N 0.000 description 1
- VUVCRYXYUUPGSB-GLLZPBPUSA-N Thr-Gln-Glu Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N)O VUVCRYXYUUPGSB-GLLZPBPUSA-N 0.000 description 1
- UDQBCBUXAQIZAK-GLLZPBPUSA-N Thr-Glu-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O UDQBCBUXAQIZAK-GLLZPBPUSA-N 0.000 description 1
- LHEZGZQRLDBSRR-WDCWCFNPSA-N Thr-Glu-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O LHEZGZQRLDBSRR-WDCWCFNPSA-N 0.000 description 1
- JKGGPMOUIAAJAA-YEPSODPASA-N Thr-Gly-Val Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)NCC(=O)N[C@@H](C(C)C)C(O)=O JKGGPMOUIAAJAA-YEPSODPASA-N 0.000 description 1
- SCSVNSNWUTYSFO-WDCWCFNPSA-N Thr-Lys-Glu Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(O)=O SCSVNSNWUTYSFO-WDCWCFNPSA-N 0.000 description 1
- SPVHQURZJCUDQC-VOAKCMCISA-N Thr-Lys-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(O)=O SPVHQURZJCUDQC-VOAKCMCISA-N 0.000 description 1
- JAJOFWABAUKAEJ-QTKMDUPCSA-N Thr-Pro-His Chemical compound C[C@H]([C@@H](C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC2=CN=CN2)C(=O)O)N)O JAJOFWABAUKAEJ-QTKMDUPCSA-N 0.000 description 1
- 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 1
- AHERARIZBPOMNU-KATARQTJSA-N Thr-Ser-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O AHERARIZBPOMNU-KATARQTJSA-N 0.000 description 1
- QYDKSNXSBXZPFK-ZJDVBMNYSA-N Thr-Thr-Arg Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O QYDKSNXSBXZPFK-ZJDVBMNYSA-N 0.000 description 1
- NHQVWACSJZJCGJ-FLBSBUHZSA-N Thr-Thr-Ile Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O NHQVWACSJZJCGJ-FLBSBUHZSA-N 0.000 description 1
- TZNNEYFZZAHLBL-BPUTZDHNSA-N Trp-Arg-Asp Chemical compound [H]N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(O)=O)C(O)=O TZNNEYFZZAHLBL-BPUTZDHNSA-N 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
- RIKLKPANMFNREP-FDARSICLSA-N Trp-Met-Ile Chemical compound C1=CC=C2C(C[C@H](N)C(=O)N[C@@H](CCSC)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O)=CNC2=C1 RIKLKPANMFNREP-FDARSICLSA-N 0.000 description 1
- MOCXXGZHHSPNEJ-AVGNSLFASA-N Tyr-Cys-Glu Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CS)C(=O)N[C@@H](CCC(O)=O)C(O)=O MOCXXGZHHSPNEJ-AVGNSLFASA-N 0.000 description 1
- IMXAAEFAIBRCQF-SIUGBPQLSA-N Tyr-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=CC=C(C=C1)O)N IMXAAEFAIBRCQF-SIUGBPQLSA-N 0.000 description 1
- NMKJPMCEKQHRPD-IRXDYDNUSA-N Tyr-Gly-Tyr Chemical compound C([C@H](N)C(=O)NCC(=O)N[C@@H](CC=1C=CC(O)=CC=1)C(O)=O)C1=CC=C(O)C=C1 NMKJPMCEKQHRPD-IRXDYDNUSA-N 0.000 description 1
- BSCBBPKDVOZICB-KKUMJFAQSA-N Tyr-Leu-Asp Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O BSCBBPKDVOZICB-KKUMJFAQSA-N 0.000 description 1
- FGVFBDZSGQTYQX-UFYCRDLUSA-N Tyr-Phe-Val Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](C(C)C)C(O)=O FGVFBDZSGQTYQX-UFYCRDLUSA-N 0.000 description 1
- PYJKETPLFITNKS-IHRRRGAJSA-N Tyr-Pro-Asn Chemical compound N[C@@H](Cc1ccc(O)cc1)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC(N)=O)C(O)=O PYJKETPLFITNKS-IHRRRGAJSA-N 0.000 description 1
- BIWVVOHTKDLRMP-ULQDDVLXSA-N Tyr-Pro-Leu Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CC(C)C)C(O)=O BIWVVOHTKDLRMP-ULQDDVLXSA-N 0.000 description 1
- GOPQNCQSXBJAII-ULQDDVLXSA-N Tyr-Val-Lys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CC1=CC=C(C=C1)O)N GOPQNCQSXBJAII-ULQDDVLXSA-N 0.000 description 1
- 108090000848 Ubiquitin Proteins 0.000 description 1
- 102000044159 Ubiquitin Human genes 0.000 description 1
- RUCNAYOMFXRIKJ-DCAQKATOSA-N Val-Ala-Lys Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCCCN RUCNAYOMFXRIKJ-DCAQKATOSA-N 0.000 description 1
- SLLKXDSRVAOREO-KZVJFYERSA-N Val-Ala-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](C)NC(=O)[C@H](C(C)C)N)O SLLKXDSRVAOREO-KZVJFYERSA-N 0.000 description 1
- VJOWWOGRNXRQMF-UVBJJODRSA-N Val-Ala-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](C)NC(=O)[C@@H](N)C(C)C)C(O)=O)=CNC2=C1 VJOWWOGRNXRQMF-UVBJJODRSA-N 0.000 description 1
- LNYOXPDEIZJDEI-NHCYSSNCSA-N Val-Asn-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](C(C)C)N LNYOXPDEIZJDEI-NHCYSSNCSA-N 0.000 description 1
- YCMXFKWYJFZFKS-LAEOZQHASA-N Val-Gln-Asp Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CC(=O)O)C(=O)O)N YCMXFKWYJFZFKS-LAEOZQHASA-N 0.000 description 1
- XWYUBUYQMOUFRQ-IFFSRLJSSA-N Val-Glu-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)O)NC(=O)[C@H](C(C)C)N)O XWYUBUYQMOUFRQ-IFFSRLJSSA-N 0.000 description 1
- 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 1
- LKUDRJSNRWVGMS-QSFUFRPTSA-N Val-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(C)C)N LKUDRJSNRWVGMS-QSFUFRPTSA-N 0.000 description 1
- VXDSPJJQUQDCKH-UKJIMTQDSA-N Val-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](C(C)C)N VXDSPJJQUQDCKH-UKJIMTQDSA-N 0.000 description 1
- DJQIUOKSNRBTSV-CYDGBPFRSA-N Val-Ile-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)O)NC(=O)[C@H](C(C)C)N DJQIUOKSNRBTSV-CYDGBPFRSA-N 0.000 description 1
- AGXGCFSECFQMKB-NHCYSSNCSA-N Val-Leu-Asp Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)[C@H](C(C)C)N AGXGCFSECFQMKB-NHCYSSNCSA-N 0.000 description 1
- RYHUIHUOYRNNIE-NRPADANISA-N Val-Ser-Gln Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N RYHUIHUOYRNNIE-NRPADANISA-N 0.000 description 1
- UJMCYJKPDFQLHX-XGEHTFHBSA-N Val-Ser-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](C(C)C)N)O UJMCYJKPDFQLHX-XGEHTFHBSA-N 0.000 description 1
- YQYFYUSYEDNLSD-YEPSODPASA-N Val-Thr-Gly Chemical compound CC(C)[C@H](N)C(=O)N[C@@H]([C@@H](C)O)C(=O)NCC(O)=O YQYFYUSYEDNLSD-YEPSODPASA-N 0.000 description 1
- JXCOEPXCBVCTRD-JYJNAYRXSA-N Val-Tyr-Arg Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)N JXCOEPXCBVCTRD-JYJNAYRXSA-N 0.000 description 1
- SSKKGOWRPNIVDW-AVGNSLFASA-N Val-Val-His Chemical compound CC(C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N SSKKGOWRPNIVDW-AVGNSLFASA-N 0.000 description 1
- 240000008042 Zea mays Species 0.000 description 1
- 235000016383 Zea mays subsp huehuetenangensis Nutrition 0.000 description 1
- 235000002017 Zea mays subsp mays Nutrition 0.000 description 1
- 230000036579 abiotic stress Effects 0.000 description 1
- 108010024078 alanyl-glycyl-serine Proteins 0.000 description 1
- 108010047495 alanylglycine Proteins 0.000 description 1
- 108010050025 alpha-glutamyltryptophan Proteins 0.000 description 1
- 238000013459 approach Methods 0.000 description 1
- 108010069926 arginyl-glycyl-serine Proteins 0.000 description 1
- 108010062796 arginyllysine Proteins 0.000 description 1
- 108010077245 asparaginyl-proline Proteins 0.000 description 1
- 108010040443 aspartyl-aspartic acid Proteins 0.000 description 1
- 108010047857 aspartylglycine Proteins 0.000 description 1
- 108010092854 aspartyllysine Proteins 0.000 description 1
- 244000052616 bacterial pathogen Species 0.000 description 1
- 230000004790 biotic stress Effects 0.000 description 1
- 239000013043 chemical agent Substances 0.000 description 1
- 238000004587 chromatography analysis Methods 0.000 description 1
- 238000003776 cleavage reaction Methods 0.000 description 1
- 238000010835 comparative analysis Methods 0.000 description 1
- 108010016616 cysteinylglycine Proteins 0.000 description 1
- 238000013461 design Methods 0.000 description 1
- 238000010586 diagram Methods 0.000 description 1
- 230000029087 digestion Effects 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
- NJDNXYGOVLYJHP-UHFFFAOYSA-L disodium;2-(3-oxido-6-oxoxanthen-9-yl)benzoate Chemical compound [Na+].[Na+].[O-]C(=O)C1=CC=CC=C1C1=C2C=CC(=O)C=C2OC2=CC([O-])=CC=C21 NJDNXYGOVLYJHP-UHFFFAOYSA-L 0.000 description 1
- 230000000694 effects Effects 0.000 description 1
- 238000001962 electrophoresis Methods 0.000 description 1
- 238000005516 engineering process Methods 0.000 description 1
- 230000010429 evolutionary process Effects 0.000 description 1
- 108010042598 glutamyl-aspartyl-glycine Proteins 0.000 description 1
- 108010057083 glutamyl-aspartyl-leucine Proteins 0.000 description 1
- 108010055341 glutamyl-glutamic acid Proteins 0.000 description 1
- 108010049041 glutamylalanine Proteins 0.000 description 1
- 108010050475 glycyl-leucyl-tyrosine Proteins 0.000 description 1
- 108010045126 glycyl-tyrosyl-glycine Proteins 0.000 description 1
- 108010050848 glycylleucine Proteins 0.000 description 1
- 108010015792 glycyllysine Proteins 0.000 description 1
- 108010037850 glycylvaline Proteins 0.000 description 1
- 230000012010 growth Effects 0.000 description 1
- 238000012165 high-throughput sequencing Methods 0.000 description 1
- 230000036039 immunity Effects 0.000 description 1
- 108010027338 isoleucylcysteine Proteins 0.000 description 1
- 230000002045 lasting effect Effects 0.000 description 1
- 108010034529 leucyl-lysine Proteins 0.000 description 1
- 108010047926 leucyl-lysyl-tyrosine Proteins 0.000 description 1
- 108010087810 leucyl-seryl-glutamyl-leucine Proteins 0.000 description 1
- 108010000761 leucylarginine Proteins 0.000 description 1
- 230000007774 longterm Effects 0.000 description 1
- 108010017391 lysylvaline Proteins 0.000 description 1
- 235000009973 maize Nutrition 0.000 description 1
- 108010016686 methionyl-alanyl-serine Proteins 0.000 description 1
- 239000000203 mixture Substances 0.000 description 1
- 230000000243 photosynthetic effect Effects 0.000 description 1
- 239000004033 plastic Substances 0.000 description 1
- 229920003023 plastic Polymers 0.000 description 1
- 238000012809 post-inoculation Methods 0.000 description 1
- 239000000843 powder Substances 0.000 description 1
- 108010077112 prolyl-proline Proteins 0.000 description 1
- 108010090894 prolylleucine Proteins 0.000 description 1
- 238000012113 quantitative test Methods 0.000 description 1
- 102000005962 receptors Human genes 0.000 description 1
- 238000011160 research Methods 0.000 description 1
- 238000010839 reverse transcription Methods 0.000 description 1
- 102220036093 rs515726130 Human genes 0.000 description 1
- 239000000523 sample Substances 0.000 description 1
- 230000007017 scission Effects 0.000 description 1
- 238000002864 sequence alignment Methods 0.000 description 1
- 108010048818 seryl-histidine Proteins 0.000 description 1
- 108010048397 seryl-lysyl-leucine Proteins 0.000 description 1
- 238000009331 sowing Methods 0.000 description 1
- 238000005507 spraying Methods 0.000 description 1
- 238000007447 staining method Methods 0.000 description 1
- 238000012360 testing method Methods 0.000 description 1
- 108010033670 threonyl-aspartyl-tyrosine Proteins 0.000 description 1
- 108010031491 threonyl-lysyl-glutamic acid Proteins 0.000 description 1
- 238000012546 transfer Methods 0.000 description 1
- 108010051110 tyrosyl-lysine Proteins 0.000 description 1
- 108010078580 tyrosylleucine Proteins 0.000 description 1
- 108010003137 tyrosyltyrosine Proteins 0.000 description 1
- 239000003643 water by type Substances 0.000 description 1
Images
Classifications
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12Q—MEASURING OR TESTING PROCESSES INVOLVING ENZYMES, NUCLEIC ACIDS OR MICROORGANISMS; COMPOSITIONS OR TEST PAPERS THEREFOR; PROCESSES OF PREPARING SUCH COMPOSITIONS; CONDITION-RESPONSIVE CONTROL IN MICROBIOLOGICAL OR ENZYMOLOGICAL PROCESSES
- C12Q1/00—Measuring or testing processes involving enzymes, nucleic acids or microorganisms; Compositions therefor; Processes of preparing such compositions
- C12Q1/68—Measuring or testing processes involving enzymes, nucleic acids or microorganisms; Compositions therefor; Processes of preparing such compositions involving nucleic acids
- C12Q1/6876—Nucleic acid products used in the analysis of nucleic acids, e.g. primers or probes
- C12Q1/6888—Nucleic acid products used in the analysis of nucleic acids, e.g. primers or probes for detection or identification of organisms
- C12Q1/6895—Nucleic acid products used in the analysis of nucleic acids, e.g. primers or probes for detection or identification of organisms for plants, fungi or algae
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12Q—MEASURING OR TESTING PROCESSES INVOLVING ENZYMES, NUCLEIC ACIDS OR MICROORGANISMS; COMPOSITIONS OR TEST PAPERS THEREFOR; PROCESSES OF PREPARING SUCH COMPOSITIONS; CONDITION-RESPONSIVE CONTROL IN MICROBIOLOGICAL OR ENZYMOLOGICAL PROCESSES
- C12Q2600/00—Oligonucleotides characterized by their use
- C12Q2600/13—Plant traits
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12Q—MEASURING OR TESTING PROCESSES INVOLVING ENZYMES, NUCLEIC ACIDS OR MICROORGANISMS; COMPOSITIONS OR TEST PAPERS THEREFOR; PROCESSES OF PREPARING SUCH COMPOSITIONS; CONDITION-RESPONSIVE CONTROL IN MICROBIOLOGICAL OR ENZYMOLOGICAL PROCESSES
- C12Q2600/00—Oligonucleotides characterized by their use
- C12Q2600/156—Polymorphic or mutational markers
Landscapes
- Chemical & Material Sciences (AREA)
- Life Sciences & Earth Sciences (AREA)
- Analytical Chemistry (AREA)
- Engineering & Computer Science (AREA)
- Organic Chemistry (AREA)
- Proteomics, Peptides & Aminoacids (AREA)
- Health & Medical Sciences (AREA)
- Biotechnology (AREA)
- Zoology (AREA)
- Wood Science & Technology (AREA)
- Immunology (AREA)
- Mycology (AREA)
- Microbiology (AREA)
- Molecular Biology (AREA)
- Botany (AREA)
- Biophysics (AREA)
- Physics & Mathematics (AREA)
- Biochemistry (AREA)
- Bioinformatics & Cheminformatics (AREA)
- General Engineering & Computer Science (AREA)
- General Health & Medical Sciences (AREA)
- Genetics & Genomics (AREA)
- Measuring Or Testing Involving Enzymes Or Micro-Organisms (AREA)
Abstract
本发明公开了小麦抗白粉病相关基因Pm41的功能型分子标记及其应用。以小麦的基因组DNA为模板，采用引物F和引物R组成的引物对扩增得到的DNA片段即为分子标记Pm41‑427。引物F为序列表的序列8所示的单链DNA分子，引物R为序列表的序列9所示的单链DNA分子。应用本发明提供的分子标记Pm41‑427，能快速筛选出具有白粉病抗性的小麦材料，从而加速小麦新品种的育成步伐。本发明具有重要的理论意义和经济价值。
Description
技术领域
本发明属于植物分子育种领域，具体涉及小麦抗白粉病相关基因Pm41的功能型分子标记及其应用。
背景技术
小麦(Triticum aestivum L.)是世界三大主粮之一，全球超过1/3的人口以其作为重要食物来源。白粉病是一种严重危害小麦生产的真菌性病害，其致病菌是禾本科专化型布氏白粉菌(Blumeria graminis f.sp.tritici)。白粉菌侵染小麦后会引起光合效率降低，从而影响植株正常生长发育，同时还会引起分蘖和成穗率降低，千粒重下降，造成减产。生产实践表明，与喷洒化学药剂相比，培育持久广谱抗病的小麦品种才是一种更加经济、环保和高效的措施。
野生二粒小麦(2n＝4x＝28，AABB)是四倍体和六倍体小麦的直接祖先，起源于中东“新月沃土”地带，广泛分布于以色列、叙利亚、黎巴嫩和土耳其等地区。野生二粒小麦经历了长期复杂的进化历程，积累了丰富的遗传多样性，对多种生物和非生物胁迫具有适应能力。研究发现野生二粒小麦蕴含有丰富的抗白粉基因，如Pm16、Pm26、Pm30、Pm36、Pm41、Pm42和Pm64等。此外，还具有与普通小麦容易杂交，杂交后代结实率高等优点，使其成为小麦抗病育种的重要遗传资源。
Pm41是一个来源于野生二粒小麦的优异抗白粉病基因，在苗期和成株期均高抗白粉病。Li(2009)等通过开发分子标记，发现6个SSR分子标记、3个EST-STS分子标记和1个ISBP标记与Pm41基因紧密连锁，将其定位在小麦3BL染色体上标记BE489472和Xwmc687之间2.7cM的遗传区间；利用中国春缺体系将Pm41定位于染色体3BL bin 0.63-1.00物理区间。Wang(2014)等利用比较基因组学方法开发了19个与Pm41紧密连锁的分子标记，将其进一步定位于标记XWGGC1505和XWGGC1507之间0.6cM的遗传区间。由于这些分子标记与Pm41基因距离较远，导致其在分子标记辅助选择育种中实用性和精确性不足。因此需要进一步开发快速、高通量、低成本的Pm41功能标记，以满足生产上利用分子标记辅助选择育种的现实需求。
发明内容
本发明的目的是提供用于鉴定待测小麦是否具有白粉病抗性的分子标记。
本发明首先保护由引物F和引物R组成的特异引物对；
所述引物F可为如下a1)或a2)：a1)序列表的序列8所示的单链DNA分子；a2)将序列8经过一个或几个核苷酸的取代和/或缺失和/或添加且与序列8具有相同功能的DNA分子；
所述引物R可为如下a5)或a6)：a5)序列表的序列9所示的单链DNA分子；a6)将序列9经过一个或几个核苷酸的取代和/或缺失和/或添加且与序列9具有相同功能的DNA分子。
所述特异引物对的功能可为如下b1)-b3)中的任一种：b1)辅助鉴定待测小麦是否具有白粉病抗性；b2)辅助筛选抗白粉病的小麦品种；b3)辅助筛选感白粉病的小麦品种。
所述特异引物对用于扩增下述与小麦白粉病抗性相关的分子标记Pm41-427。
本发明还保护一种试剂盒，可包括所述特异引物对。
所述试剂盒还可包括限制性内切酶HindⅢ。
所述试剂盒中还可包括用于PCR扩增的常规试剂和/或用于基因组提取的常规试剂和/或用于琼脂糖凝胶电泳的常规试剂和/或用于限制性内切酶酶切的常规试剂。
所述试剂盒的功能可为如下b1)-b3)中的任一种：b1)辅助鉴定待测小麦是否具有白粉病抗性；b2)辅助筛选抗白粉病的小麦品种；b3)辅助筛选感白粉病的小麦品种。
本发明还保护上述任一所述特异引物对或上述任一所述试剂盒的应用，可为如下b1)-b3)中的任一种：b1)辅助鉴定待测小麦是否具有白粉病抗性；b2)辅助筛选抗白粉病的小麦品种；b3)辅助筛选感白粉病的小麦品种。
以待测小麦的基因组DNA为模板，采用所述特异引物对扩增得到的DNA片段即为分子标记Pm41-427。
本发明还保护以待测小麦的基因组DNA为模板，采用所述特异引物对扩增得到的DNA片段(即本发明保护与小麦白粉病抗性相关的分子标记Pm41-427)。
本发明还保护所述分子标记Pm41-427的应用，可为如下b1)-b3)中的任一种：b1)辅助鉴定待测小麦是否具有白粉病抗性；b2)辅助筛选抗白粉病的小麦品种；b3)辅助筛选感白粉病的小麦品种。
应用所述特异引物对、所述试剂盒或所述分子标记Pm41-427辅助鉴定待测小麦是否具有白粉病抗性的方法可如下：以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，如果酶切产物中具有189bp的DNA片段，则待测小麦具有或疑似具有白粉病抗性，否则待测小麦不具有或疑似不具有白粉病抗性。
应用所述特异引物对、所述试剂盒或所述分子标记Pm41-427辅助筛选抗白粉病的小麦品种的方法可如下：以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，如果酶切产物中具有189bp的DNA片段，则待测小麦为候选的抗白粉病的小麦品种；否则待测小麦不为候选的抗白粉病的小麦品种。
应用所述特异引物对、所述试剂盒或所述分子标记Pm41-427辅助筛选感白粉病的小麦品种的方法可如下：以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，如果酶切产物中“具有214bp的DNA片段且不具有189bp的DNA片段”或“不具有任何DNA片段”，则待测小麦为候选的感白粉病的小麦品种；否则待测小麦不为候选的感白粉病的小麦品种。
本发明还保护检测待测小麦是否具有白粉病抗性的方法，可为方法R1)、方法R2)、方法S1)、方法S2)、方法T1)和方法T2)中的至少一种。
方法R1)检测待测小麦的基因组中是否含有序列表中序列4所示的DNA片段和序列表中的序列5所示的DNA片段，如果待测小麦的基因组中含有序列表中序列4所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性。
方法R2)检测待测小麦的基因组中是否含有序列表中序列4所示的DNA片段和序列表中的序列5所示的DNA片段，然后进行如下判断：如果待测小麦的基因组中含有序列表中序列4所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果待测小麦的基因组中含有序列表中序列5所示的DNA片段且不含有序列表中序列4所示的DNA片段，则待测小麦不具有或疑似不具有白粉病抗性。
方法S1)以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，如果酶切产物中具有189bp的DNA片段，则待测小麦具有或疑似具有白粉病抗性。
方法S2)以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，然后进行如下判断：如果酶切产物中具有189bp的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果酶切产物中“具有214bp的DNA片段且不具有189bp的DNA片段”或“不具有任何DNA片段”，则待测小麦不具有或疑似不具有白粉病抗性。
方法T1)以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；将所述PCR扩增产物进行测序，然后进行如下判断：如果PCR扩增产物含有序列表中序列6所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性。
方法T2)以待测小麦的基因组DNA为模板，采用所述特异引物对进行PCR扩增，得到PCR扩增产物；将所述PCR扩增产物进行测序，然后进行如下判断：如果PCR扩增产物含有序列表中序列6所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果PCR扩增产物“含有序列表中序列7所示的DNA片段且不含有序列表中序列6所示的DNA片段”或“不含有任何DNA片段”，则待测小麦不具有或疑似不具有白粉病抗性。
上述任一所述189bp的DNA片段的核苷酸序列如序列表中序列6自5’末端起第1至189位所示。
上述任一所述214bp的DNA片段的核苷酸序列如序列表中序列6所示。
上述任一所述白粉病可为白粉菌引起的病害。所述白粉菌具体可为白粉菌生理小种E09。
实验证明，应用本发明提供的分子标记Pm41-427，能快速筛选出具有白粉病抗性的小麦材料，从而加速小麦新品种的育成步伐。本发明具有重要的理论意义和经济价值。
附图说明
图1为检测TZ-2、Langdon和“TZ-2和Langdon的杂交F1代”对白粉菌的抗性。
图2为Pm41基因的图位克隆。
图3为Pm41基因精细定位图。
图4为CNL基因的表达分析。
图5为CNL基因互补和过表达载体示意图及其转基因T2代家系的分子鉴定和白粉病抗性鉴定。
图6为分子标记Pm41-427的多态性检测。
图7为分子标记Pm41-427对42份野生二粒小麦的部分检测结果。
图8为分子标记Pm41-427对262份六倍体中国小麦微核心种质的部分检测结果。
具体实施方式
以下的实施例便于更好地理解本发明，但并不限定本发明。下述实施例中的实验方法，如无特殊说明，均为常规方法。下述实施例中所用的试验材料，如无特殊说明，均为自常规生化试剂商店购买得到的。以下实施例中的定量试验，均设置三次重复实验，结果取平均值。
下述实施例中，小麦苗期白粉病抗性鉴定方法具体参考如下文献：liu ZY，SunQX，Ni ZF，Yang TM(1999)Development of SCArmarkers linked to the Pm21geneconferring resistance to powdery mildew in common wheat.Plant Breeding 118：215-219。
抗白粉病亲本TZ-2、感白粉病亲本Langdon和白粉菌生理小种E09均记载于如下文献中：Wang ZZ，Cui Y，Chen YX(2014)Comparative genetic mapping and genomicregion collinearity analysis of the powdery mildew resistancegenePm41.Theoretical and Applied Genetics，127:1741-1751。文献中的IW2与本文中的抗白粉病亲本TZ-2是同一材料。在下文中，抗白粉病亲本TZ-2简称TZ-2，感白粉病亲本Langdon简称Langdon，白粉菌生理小种E09为简称白粉菌。
普通小麦品种Fielder为转基因受体材料。普通小麦品种Fielder为高感白粉病的小麦品种。在下文中，普通小麦品种Fielder简称Fielder。
pCAMBIA1300质粒为构建互补转基因载体的基础载体，购买于生化试剂公司，采用限制性内切酶BamHⅠ和HindⅢ进行双酶切，然后利用同源重组方法构建互补转基因载体。
pTPCK303质粒为构建过表达转基因载体的基础载体，购买于生化试剂公司，采用限制性内切酶KpnⅠ和SpeⅠ进行双酶切，然后利用同源重组方法构建过表达转基因载体。
下述实施例中涉及的引物的名称及其核苷酸序列详见表1。
表1
实施例1、小麦抗白粉病基因Pm41(以下简称Pm41基因)高密度遗传连锁图谱构建
1、以TZ-2为父本，Langdon为母本进行杂交，得到杂交F1代。
2、完成步骤1后，采用小麦苗期白粉病抗性鉴定方法检测TZ-2、Langdon或杂交F1代的白粉病抗性；采用考马斯亮蓝染色方法(记载于如下文献中：李健强，刘西莉和王红梅(2002)荧光素钠和考马斯亮蓝应用于小麦白粉病菌染色效果比较.菌物系统，21(4):592-596)检测TZ-2、Langdon或杂交F1代的白粉病抗性。
检测结果见图1(F1为杂交F1代，A为叶片接种白粉菌反应，B为白粉菌孢子染色)。结果表明，TZ-2和杂交F1代均抗白粉病，Langdon则高感白粉病。
3、取步骤1得到的杂交F1代，播种于大田，通过自交方式，得到2448个F2分离群体。
4、完成步骤3后，采用小麦苗期白粉病抗性鉴定方法检测来自于F2分离群体的F2:3家系的白粉病抗性。
5、完成步骤4后，从F2:3家系中挑选90个家系(包括45个纯合抗病家系和45个纯合感病家系)，分别在苗期接种白粉菌。接种后24h，将每个抗病家系采集等量叶片混合得到抗病混池，每个感病家系采集等量叶片混合得到感病混池，然后分别提取总RNA，构建转录组测序文库。
6、完成步骤5后，采用高通量测序平台Illumina HiSeq 2000分别对抗病混池和感病混池进行转录组双末端测序。测序原始数据采用小麦BSR-Seq基因定位技术进行分析。
7、完成步骤6后，根据分析结果开发了7个与Pm41紧密连锁的分子标记(见图2中B，相应的引物见表1中精细定位引物)。
8、完成步骤7后，利用文献(Wang ZZ，Cui Y，Chen YX(2014)Comparative geneticmapping and genomic region collinearity analysis of the powdery mildewresistance gene Pm41.Theoretical and Applied Genetics，127:1741-1751)报道的与Pm41基因紧密连锁且位于基因两侧的SSR标记Barc84和Barc77(引物序列见表1)筛选2448个F2分离群体，鉴定出交换单株。
9、完成步骤8后，采用新开发的7个与Pm41紧密连锁分子标记检测步骤8筛选得到的交换单株。用Mapmaker 3.0计算分子标记与Pm41基因的遗传距离。用Mapdraw V2.1软件绘制遗传连锁图谱。最终Pm41基因被定位于分子标记M171和M160之间0.08cM的遗传区间，其中分子标记M78和M97与Pm41基因共分离(见图2中B和图3)。
实施例2、Pm41基因物理图谱构建
1、构建抗白粉病亲本TZ-2的BAC文库(记载于如下文献中：梁永.野生二粒小麦抗白粉病基因MlIW170物理图谱构建和粗山羊草2DS同源区域比较分析[D]；中国农业大学；2015年)。构建的抗白粉病亲本TZ-2的BAC文库共包含326,784个克隆，平均插入片段长度为120kb，约3×基因组覆盖度，含有HindIII酶切位点。BAC文库的阳性克隆被分装在851个384孔板中。将每个384孔板中的所有BAC克隆混合，混合后摇菌提取BAC质粒，建立一级筛选混池，共851个。将851个混池编号后，放置于10×10的100孔盒中，每个100孔盒中的每一个横排10个混合，每一个列池10个混合，混合后摇菌提取BAC质粒，建立了二级筛选混池，共176个。
2、完成步骤1后，在Pm41精细定位的基础上，利用定位区间两侧的分子标记开发特异性探针筛选抗白粉病亲本TZ-2的BAC文库，得到9个BAC阳性克隆(见图2中C，9个BAC阳性克隆分别为412O18、783K1、612H21、391A10、614D21、318M1、111C12、446G18和599A11)。
3、完成步骤2后，利用QIAGEN Large-Construct Kit试剂盒提取BAC阳性克隆的质粒，然后在Illumina Hi Seq2500PE250测序平台进行测序，拼接组装获得了2条Contigs，大小分别为140kb和350kb。利用TriAnnot平台进行基因注释(Leroy P，Guilhot N，Sakai H，et al(2012)TriAnnot：a versatile and high performance pipeline for theautomated annotation of plant genomes.Frontiers in Plant Science，3)，共获得6个基因，分别是TzRlpA基因(编码稀有脂蛋白酶A)、TzGNT基因(编码糖基转移酶)、TzCNL基因、TzHP1基因(编码假定蛋白1)、TzHP2基因(编码假定蛋白2)和TzHP3基因(编码假定蛋白3)(见图2中D)。
实施例3、CNL基因的表达分析
hpi(hour post inoculation)表示接种后的时间。
检测CNL基因或cnl基因的引物为表1中所示的41R-315。检测Actin基因的引物为表1中所示的Actin。
1、待TZ-2或Langdon长至2片叶时，利用白粉菌进行拂弹接种(记载于如下文献中：liu ZY，Sun QX，Ni ZF，Yang TM(1999)Development of SCArmarkers linked to thePm21gene conferring resistance to powdery mildew in common wheat.PlantBreed118:215-219)。
2、完成步骤1后，在0hpi、2hpi、4hpi、12hpi、18hpi、24hpi、36hpi、48hpi、60hpi和72hpi时间点分别取叶片，然后提取RNA，反转录，得到cDNA。
3、分别以步骤2得到的cDNA为模板，进行半定量检测。
半定量检测结果见图4中A(CNLTZ-2表示TZ-2中CNL基因，cnlLangdon表示Langdon中cnl基因，TaActin表示小麦内参Actin基因)。结果表明，接种白粉菌后10个时间点，TZ-2中能检测到CNL基因表达，Langdon中则一直不能检测到CNL基因表达。
4、分别以步骤2得到的cDNA为模板，进行实时荧光定量PCR检测。
实时荧光定量PCR检测结果见图4中B。结果表明，4hpi时CNL基因表达明显上调，12hpi时CNL基因的表达量达到最高峰；随后，CNL基因表达表现下调。因此，TZ-2中CNL基因能够受到白粉菌强烈的诱导表达。
实施例4、CNL基因的克隆
1、采用CNL-1(引物序列见表1)从BAC阳性克隆391A10的质粒中扩增CNL基因，测序拼接获得一个11730bp的核苷酸序列(如序列表中序列1所示)，基因结构预测发现包含有一个完整的CC-NBS-LRR类型的基因(见图2中E)。
2、取TZ-2接种白粉菌后24h的叶片，提取总RNA。
3、完成步骤2后，以TZ-2的总RNA反转录后cDNA为模板，采用CNL-2(引物序列见表1)进行PCR扩增。同时采用
RACE 5'/3'Kit RACE试剂盒对CNL基因的5'和3'末端进行扩增。最终获得一个3585bp的核苷酸序列。
4、将步骤3得到的核苷酸序列(3585bp)与步骤1得到的核苷酸序列(11730bp)进行比对。结果发现CNL基因具有两个外显子，11730bp的核苷酸序列包含基因编码区(序列表中序列1自5’末端起第5379-6206位和第6622-8748所示)、启动子区(序列表中序列1自5’末端起第1-5043位所示)、5'-UTR区(序列表中序列1自5’末端起第5044-5378位所示)和3'-UTR区(序列表中序列1自5’末端起第8749-9135位所示)，全长ORF区的核苷酸序列为2955bp。
CNL基因的编码区(2955bp)的核苷酸序列如序列表中序列2所示，编码序列表中序列3所示的CNL蛋白。
实施例5、CNL互补转基因T2家系和CNL基因过表达T2家系的获得与白粉病抗性鉴定
一、重组质粒的构建
1、重组质粒pCAMBIA1300-CNL的构建
(1)取pCAMBIA1300质粒，用限制性内切酶BamHⅠ和HindⅢ进行双酶切，回收约8900bp的载体骨架。
(2)以序列表中序列1所示的DNA片段为模板，采用41HB-1(见表1)进行PCR扩增，得到PCR扩增产物。
(3)利用pEASY-Uni Seamless Cloning and Assembly Kit(CU101-01)同源重组试剂盒将步骤(2)获得的PCR扩增产物和步骤(1)回收的载体骨架进行同源重组，得到重组质粒pCAMBIA1300-CNL(见图5中A)。重组质粒pCAMBIA1300-CNL中，CNL基因由其自身启动子(序列表中序列1自5’末端起第1-5043位所示)驱动。
2、重组质粒pTPCK303-CNL的构建
(1)取pTPCK303质粒，用限制性内切酶Kpn1和Spe1进行双酶切，回收约14000bp载体骨架。
(2)以序列表中序列2所示的DNA片段为模板，采用41OE-1(见表1)进行PCR扩增，得到扩增产物。
(3)利用pEASY-Uni Seamless Cloning and Assembly Kit(CU101-01)同源重组试剂盒将步骤(2)获得的PCR扩增产物和步骤(1)回收的载体骨架进行同源重组，得到重组质粒pTPCK303-CNL。重组质粒pTPCK303-CNL中，CNL基因由玉米的Ubiquitin启动子驱动(见图5中B)。
二、重组农杆菌的获得
1、采用热激转化法，将重组质粒pCAMBIA1300-CNL导入根癌农杆菌EHA105，得到重组农杆菌，命名为EHA105/pCAMBIA1300-CNL。
2、采用热激转化法，将重组质粒pTPCK303-CNL导入根癌农杆菌EHA105，得到重组农杆菌，命名为EHA105/pTPCK303-CNL。
三、转CNL基因小麦的获得
1、采用农杆菌介导的遗传转化方法，将EHA105/pCAMBIA1300-CNL转化高感白粉病的普通小麦材料Fielder，然后自交2代，得到5个CNL互补转基因T2家系，依次命名为T2CNL-1至T2CNL-5。
2、采用农杆菌介导的遗传转化方法，将EHA105/pTPCK303-CNL转化高感白粉病的普通小麦材料Fielder，然后自交2代，得到5个CNL基因过表达T2家系，依次命名为T2OECNL-1至T2OECNL-5。
四、分子鉴定
1、提取小麦叶片的基因组DNA。
小麦为TZ-2、Fielder、T2CNL-1、T2CNL-2、T2CNL-3、T2CNL-4、T2CNL-5、T2OECNL-1、T2OECNL-2、T2OECNL-3、T2OECNL-4或T2OECNL-5。
2、以小麦叶片的基因组DNA为模板，采用引物进行PCR扩增(其中T2CNL-1、T2CNL-2、T2CNL-3、T2CNL-4和T2CNL-5均用41R-483(见表1)进行PCR扩增，T2OECNL-1、T2OECNL-2、T2OECNL-3、T2OECNL-4和T2OECNL-5均用41R-476(见表1)进行PCR扩增，TZ-2和Fielder分别用41R-483和41R-476进行PCR扩增)，得到PCR扩增产物，然后进行如下判断：
如果某小麦采用41R-483进行PCR扩增，得到的PCR扩增产物中含有约420bp的DNA片段，则该小麦含有CNL基因；否则该小麦不含有CNL基因。
如果某小麦采用41R-476进行PCR扩增，得到的PCR扩增产物中含有约750bp的DNA片段，则该小麦含有CNL基因；否则该小麦不含有CNL基因。
实验结果见图5中C和D(“+”表示阳性，即含有CNL基因，“-”表示阴性，即不含CNL基因)。结果表明，5个CNL互补转基因T2家系和5个CNL基因过表达T2家系也均为阳性。
五、白粉病抗性鉴定
采用小麦苗期白粉病抗性鉴定方法检测小麦的白粉病抗性。小麦为TZ-2、Fielder、T2CNL-1、T2CNL-2、T2CNL-3、T2CNL-4、T2CNL-5、T2OECNL-1、T2OECNL-2、T2OECNL-3、T2OECNL-4或T2OECNL-5。
检测结果见图5中C和D。结果表明，5个CNL互补转基因T2家系和5个CNL基因过表达T2家系均表现为高抗或免疫白粉病。
上述结果表明，CNL基因在普通六倍体小麦中具有良好的抗白粉病功能，同时也证实CNL基因就是Pm41基因。
实施例6、小麦Pm41基因分子标记的开发和多态性检测
一、CNL基因与cnl基因的核苷酸序列比对
本发明的发明人从TZ-2中扩增获得CNL基因，从Langdon中扩增获得cnl基因。将CNL基因与cnl基因进行比对，结果发现CNL基因与cnl基因的编码区存在14处SNPs差异，分别为A18G、G22A、G90T、A108C、G226T、C231T、C270A、A365G、T498G、G549T、A765G、G922A、A2304C、G2903C和G3249C(数字前的碱基是CNL基因的碱基，数字后的碱基是cnl基因的碱基，数字是从基因起始密码子(计为0)开始的碱基顺序位置)。
二、分子标记Pm41-427的开发
根据TZ-2中的CNL基因和Langdon中的cnl基因在起始密码子开始第22位的差异G22A开发dCAPS标记Pm41-427(简称分子标记Pm41-427)，限制性内切酶为HindⅢ。dCAPS标记开发方法参考Neff MM，Turk E and Kalishman M(2002)Web-based primer design forsingle nucleotide polymorphism analysis.Trends in Genetics，18:613-615.
扩增分子标记Pm41-427的核苷酸序列如下所示：
引物F：5’-CGGGTACATCTGACTGTTCAATTTACA-3’(序列表中序列8)；
引物R：5’-GAGGGAGTTCATCACCCCAGTGGCAAAGCT-3’(序列表中序列9)。
三、多态性检测
限制性内切酶HindⅢ为Takara公司的产品。2×PCR Mix为北京擎科新业生物技术有限公司的产品。10×Buffer为限制性内切酶HindⅢ中的组件。
待测小麦为TZ-2、Langdon或实施例1中步骤3的F2分离群体。
1、采用植物DNA快速提取试剂盒提取待测小麦叶片的基因组DNA。
2、以小麦叶片的基因组DNA为模板，用步骤二中引物F和引物R组成的引物对进行PCR扩增，得到PCR扩增产物。
PCR反应体系(10μL)：待测小麦叶片的基因组DNA(25ng/μL)2μL、2×PCR Mix5μL、引物F的水溶液(浓度为10μmol/L)1μL、引物R的水溶液(浓度为10μmol/L)1μL和ddH2O 1μL。
PCR反应条件：94℃5min；94℃30s，58℃30s，72℃30s，35个循环；72℃7min。
3、取步骤2得到的PCR扩增产物，用限制性内切酶HindⅢ酶切，得到酶切产物；然后将酶切产物进行8％琼脂糖凝胶电泳，根据电泳结果进行如下判断：如果酶切产物为带型A(只显示一条带，为189bp)，则待测小麦的基因型为基因型I；如果酶切产物为带型B(显示两条带，为189bp和214bp)，则待测小麦的基因型为基因型II；如果酶切产物为带型C(只显示一条带，为214bp)，则待测小麦的基因型为基因型III；如果扩增产物和酶切产物不显示条带，则待测小麦的基因型为基因型IV。
酶切体系(10μL)：PCR扩增产物5μL、限制性内切酶HindⅢ1μL、10×Buffer1μL和ddH2O 3μL。
酶切反应条件：37℃30min。
部分实验结果见图6。TZ-2的基因型为基因型I。Langdon的基因型为基因型III。F2分离群体的基因型有三种，一种为基因型I(与TZ-2的基因型和白粉病抗性相同，即图6中的纯合抗病单株)，一种为基因型III(与Langdon的基因型和白粉病抗性相同，即图6中的纯合感病单株)，另一种为基因型II(高抗白粉病，即图6中的杂合单株)。
上述结果表明，步骤二开发的分子标记Pm41-427具有较高的多态性。
实施例7、分子标记Pm41-427的应用
一、应用一
42份野生二粒小麦由以色列海法大学进化研究所Nevo教授惠赠。42份野生二粒小麦具体记载于如下文献中：解超杰，孙其信，杨作民(2003)以色列野生二粒小麦苗期抗病性鉴定.麦类作物学报，23(2):39-42。42份野生二粒小麦的具体名称见表2中第1列。
1、按照实施例6中步骤三的方法分别对42份野生二粒小麦、Langdon和TZ-2进行基因分型。
部分实验结果见图7和表2中第2列。结果表明，IW1、IW2、IW3、IW7、IW8和IW10的基因型为基因型I，其余36份野生二粒小麦的基因型为基因型III。
表2
| 材料名称 | 基因型 | 白粉病抗性 | 材料名称 | 基因型 | 白粉病抗性 |
| TZ-2 | I | 抗病 | IW40 | III | 感病 |
| Langdon | III | 感病 | IW41 | III | 感病 |
| IW1 | I | 抗病 | IW42 | III | 抗病 |
| IW2 | I | 抗病 | IW43 | III | 感病 |
| IW3 | I | 抗病 | IW44 | III | 感病 |
| IW4 | III | 抗病 | IW45 | III | 感病 |
| IW5 | III | 抗病 | IW46 | III | 感病 |
| IW6 | III | 抗病 | IW47 | III | 抗病 |
| IW7 | I | 抗病 | IW48 | III | 抗病 |
| IW8 | I | 抗病 | IW50 | III | 感病 |
| IW9 | III | 感病 | IW51 | III | 感病 |
| IW10 | I | 抗病 | IW52 | III | 感病 |
| IW25 | III | 感病 | IW53 | III | 感病 |
| IW31 | III | 感病 | IW54 | III | 感病 |
| IW32 | III | 感病 | IW55 | III | 感病 |
| IW33 | III | 感病 | IW57 | III | 感病 |
| IW34 | III | 感病 | IW58 | III | 感病 |
| IW35 | III | 感病 | IW59 | III | 感病 |
| IW36 | III | 感病 | IW60 | III | 感病 |
| IW37 | III | 感病 | IW61 | III | 感病 |
| IW38 | III | 感病 | IW62 | III | 感病 |
| IW39 | III | 感病 | IW73 | III | 感病 |
2、采用小麦苗期白粉病抗性鉴定方法分别检测42份野生二粒小麦的白粉病抗性。
实验结果见表2中第3列。结果表明，基因型I的野生二粒小麦(即IW1、IW2、IW3、IW7、IW8和IW10)均表现为免疫或高抗白粉病；36份基因型III的野生二粒小麦中，30份野生二粒小麦表现为感白粉病，6份野生二粒小麦表现高抗白粉病。
上述结果表明，野生二粒小麦中，基因型为基因型I的小麦为免疫或高抗白粉病，基因型为基因型III的小麦为或疑似为感白粉病。
二、应用二
262份六倍体中国小麦微核心种质由中国农业科学院作物科学研究所张学勇研究员提供，具体记载于如下文献中：Dong YC，Cao YS，Zhang XY，Liu SC，Wang LF，You GX，Pang BS，Li LH，Jia JZ(2003)Establishment of candidate core collections inChinese common wheat germplasm.Journal of Plant Genetic Resources，4(1):1-8.和Hao CY，Dong YC，Wang LF，You GX，Zhang HN，Ge HM，Jia JZ，Zhang XY(2008)Geneticdiversity and construction of core collection in Chinese wheat geneticresources.Chinese Science Bulletin，53(8):908-915。
1、按照实施例6中步骤三的方法分别对262份六倍体中国小麦微核心种质进行基因分型。
实验结果见图8和表3中第2列和第5列。结果表明，185份六倍体中国小麦微核心种质的基因型为基因型III；77份六倍体中国小麦微核心种质的基因型为基因型IV。
表3
2、采用小麦苗期白粉病抗性鉴定方法分别检测262份六倍体中国小麦微核心种质的白粉病抗性。
实验结果见表3中第3列和第6列。结果表明，185份基因型III的六倍体中国小麦微核心种质中，174份六倍体中国小麦微核心种质表现为感白粉病，11份六倍体中国小麦微核心种质表现高抗白粉病。77份基因型IV的六倍体中国小麦微核心种质中，76份六倍体中国小麦微核心种质表现为感白粉病，1份六倍体中国小麦微核心种质表现高抗白粉病。
上述结果表明，六倍体中国小麦微核心种质中，基因型为基因型III的小麦为或疑似为感白粉病。
上述结果表明，检测待测小麦的基因型，基因型I或基因型II的小麦为抗白粉病，基因型III或基因型IV的小麦为或疑似为感白粉病。因此，实施例6开发的分子标记Pm41-427可用于辅助育种。
<110> 中国科学院遗传与发育生物学研究所
<120> 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用
<160> 9
<170> PatentIn version 3.5
<210> 1
<211> 11730
<212> DNA
<213> Artificial sequence
<400> 1
gttcctgatt atttagtata catgtatggt aagtataagt agtttaacat gaccaatcca 60
tggcatcata tgtttagagg atatgatatg ttgcattgtt gttgtcattg agtgttttca 120
aatcccgaca aatcttctca cgaccgtcat ctccctccga actagatatg gagattccgg 180
acaaaattgt cccatgctcg tcatcttcct tgagtccggt ggtcggcggc atgtgggaag 240
ctcgcatcca tggccgatgg agaggctcga gctcgtgggc gaaggttggg aagaggggcg 300
gcggaactgc ttggtcaaca ccgaggatgg aggaagaggg cgacgggagc catggccgac 360
caggacggcg tgcgaggtcg gaggggcggc ggaactgctt ggttagcacc gatgatggag 420
gaagagagcg acgggagcca tggccgacta ggagcagcgt gcaaggtcag aggggtggct 480
cgccagccgg ggcttgccgc tagggtaagg acggtgctag ctgcagcctg gcagggatgg 540
ccgtcgcacg gaagtagggg tcgcgactgg tgctagtgtg ttgtcacccg atgacccggc 600
tgtaggtaca accaggtttg catacaattt cagaattcat ttcagaaagt aacaaaaact 660
tggctagttt tatattacag ctcacagaaa cctcatgtga cccttacttt caaactacta 720
ggttatactt ttgccttgaa acacaagtac agatgggatg ggttgatgag acggtcgatc 780
tatgaatctc acattttgat cttgagatgg ggtgatctca acatctttca tcttagcggc 840
cagtccatgg agggttccta cacgtcagag taggtccaac ataatcatgt catttttgtt 900
tcgacaaaaa acatagagat taaattgtgt aaattaacaa ccaaaccaag ttcgttttga 960
gatcttttat attaggaacg tgttgttaca tcatggatca acaattgtgt atctttgttt 1020
actccctagt tgccattacg aaattcaaac ttgggatatg tgtcaaatat ttgtgccctt 1080
gccggtcaga tccgtcctgc accggaacgc gcacgagggg gcaaaggagc cctgccgccg 1140
tcacgccctc tggcggtgac agaggagggg aggagcggga gggagggcgc tcggaggcgt 1200
caacagatcc ctctcgtcgc cgtccgcagc tcgcgccacc tggtgcttgg gaccacagat 1260
ctgggcgaca tcaccgccgc caccatgacc caccacatcc caccatcgaa ggtcacccgc 1320
ctcccgacgt agccgctagc aagccaaatc gcgcatccgc gaccctccgc atagtgacgc 1380
caccttgccc gtcagatccg tcccgcatca gatccacgcg cgagggggcg aagaagccct 1440
gtcgccgccc ctaggcggac ttcacccagt catgccctct ggcggcagca agggagggga 1500
ggagggggag gcggctaggg attcgtcccg tcgcctgctg gggcgactct ggagggagtg 1560
atctcaacat ctttcatctt aggaaaaagt ccattttaaa ccttgaactt gtagatattc 1620
ggcgaaatga tcccccacgt cgaaatccct gacaccttga actatgcaat cacggtctaa 1680
atcaaccctg acagatgtca accgggattc acctgactag tgggctagaa agcgacagtc 1740
ttgaccacgg cgcgctgggc cggcccattt ttttgaagtt acagtgcgaa aaaaaatatc 1800
tatgcgtgaa actttttctt ttttatataa tccgtgaaaa aatatctaat gcatgatttt 1860
tttatataat ccgtgaaaaa atatctaatg tgtgaaactt ttttcaactt gtttttaaaa 1920
tttcgaacct tttgaaaaat ctggtggaca ttttaaaaat cattgaactt tttctaaatc 1980
catgtttttc cacaattcat caactttttt tcgaaattga tgaacttttt ccaaacctgt 2040
gaactgttat tgaaaattta tgagtttttt taaatcggtg tttttaaaaa aatatgaact 2100
ttcatccgaa gtcttcccca cgctgagcag gcccattttt ctattacatt gctataaaat 2160
atctaatgca ggaaactttt tcctttttta tataacccgt gaaaaaatat ttaattcatg 2220
tttttccttt ttcatataat acgtaaaaaa tatctaatgc gtgacactat tttaaatatg 2280
tttttcaatt tgtgaacttt tttaccaagc tcgtgaattt ttagaatctc tgaaattttt 2340
ttgaatccat gaacattttt ctatatagat gagaaaaatt ccaaacctgt gaactttttt 2400
ttgaatattg atgggatttt tcgaaaatcg ttgaatttta catttatgta atttaaaccg 2460
gagcgttgac cacggtgtcg tgattttttc gccatattca tgatcttttt ttattttcgt 2520
gaactatttg gagttttttc catttccgtg aactaaagaa gggtcaaata gtactaattt 2580
gcagcagcat acaaataagt gaagtttacg tagtggttaa tggctatagc tgttaacgta 2640
ctgcactgca gttcgactat tgctactaac atattaattg ctgagtcttt ttgttttctt 2700
ttgcgttctc tgtaggttgc tgggcttcac agactgcacg acgtggatag cctgctgggc 2760
cagactgggc tcgcctagtt tttttctttt ggcgcgctac caatcctggg cacggtacac 2820
aatccccgtg ttgcaaatga ccctacggtt cgttttagac cgggattgca tagtttaaag 2880
tacaattgac agggatttga tgttggggtt tgtttcgcgt aacctctaca aaaatcttag 2940
agattcaacc atgtaaatta tcaaccaaac caatttagtt ttgagattta cttattggcg 3000
tatcacagat ttgtcaaaaa caacctattc tttaatttac tataaaaagg aaaaaaaatg 3060
tatgcatacc taggtgtgtg gttccagatg acttggcgat aagtgggcgt gttactcttg 3120
tattccattc acatgtgact tttgtaggat cctacaacgt gctcagtgag acgatgggcg 3180
tgtgagaatt tgttgttgga agaacttcat aaattagaat aatatatttt ttcattgttt 3240
tacttgtaat tgcaacgtca caccgatgtt gcaatatatt tccgaatata accacaaata 3300
gttattctgt tgaaacacgt aatctctatt acaacaaaat gtctgcccgc gcaacggaat 3360
ttgtttgtca caatatcgct accatattgc aacgccgatg cggtttgttg caataggtag 3420
ccactttttt gcaacaatat ttgtagctaa tgcaacgaca agttgatgtt gcaatatctt 3480
tccgaatatt gccacaaata gttattccgt tgaaatacct agtctctatt acaacaaaat 3540
gactaccctc acgcaacgga atttgtttct cgcaatatcg ctaccatatt gcaacgtcga 3600
cacggtttgt tgcaatatga ggcctgtatt gcaacaaagt tagtatttat cgtgaccaaa 3660
ctaagttatg gcagtatgcg cagcctatta ccacaatcgg agaatttgtg gtaatattgt 3720
attttattgc cacaaaatgt agttgtagca ataaattttg ttgcaataga ccggaatctt 3780
tgtagtgcga gtctgtgaag gtttggaagt cgcctgaaga cttaccacga gtgattggac 3840
gaggtctgtg tgaccttagt tcaaggagaa tacggtgagg acttggtgtc ctgggctgcg 3900
tgctcagcga ttgggtgtcc gggactgtgt gtcctcgagt ttaaatactc agccgctcca 3960
accagacgta caactgagac agcagttgga actggtctac caaatcattg tcttcaccaa 4020
ccttactggt tctattccct caactctttc attttctcat cactgtgttg agtgattgtt 4080
tatatctgtg tttgaagact ttgactgaag actttctcaa tttcctcagt tcaatttctt 4140
cagtctgttt gtcttcatct tgtgttatcc tgtgtttacg ctttctgtgc tctgtgcatg 4200
tcttcatttc atcatgatga ccatgcctgt attctgttat gcttacttct gagtacttat 4260
tccgctgcaa gtagttcttt gctaaggaat ttcctcaccg gtaaattctt cataaaatcg 4320
cctattcact cccctctagt tgatataatg cactttcaga gcgcccgctt ttgtccgcac 4380
ggacccattt gtggcccatt tttgcgttca ctttgggccg ggagggcgga caaacggaca 4440
gcaggcggtc acggacgtcc gtttggatcg ccccgttggg ccaacttttg tgtccggatg 4500
acctaaacgg acaaaatgag tcgccccatt ggagttgctc ttataacaaa ttcggtcgtc 4560
ggttgtagca tggccgtgca gcggcggagc tgccactacg ccatgaccgt ggccgcagga 4620
ggacaagaag gtaagtggtg gacgaccatg gcgcccgccc aaggagctcg tcggtgtcga 4680
cctcccgttc atgctacaag gagtccgccc caaggagctc acaggtgaca accatggcgc 4740
ccgcgcaagg agctcgccgg tgacgaccat ggcgcccgca caaagagctc gccgacgtcg 4800
cacgagataa ggaaaagaag agagaaggta tgagcgagct tgtgttttta taggccaggg 4860
gaatgaatgg acgagaggag tccagaagag gataagacat cgtagaggat aaggcagcaa 4920
gccacgcgac cggtcgaaag tttggttacc atcaattatt ggtgcacttg ccttggaaaa 4980
ttcaccccag tgcactagag aagataagac caatcagtgc aaccgctccc ctcttcttca 5040
cccgcctagc tttactcaac gcctctggct gcgccagcga caacatcaag ggagcaacag 5100
attcgatcgg cagccaggct gagcaatggc cgccgagcat cagtagaggt acgttatgac 5160
catgccacga atttagctct tgattcccca tgagccatga gattagttag atcgggtaca 5220
tctgactgtt caatttacag tcttctttct cagatctgaa cctactctgc tcttggtagc 5280
agcagcttgt acttcttctc aagtcaccac ggcagtatct acactcttag attctgtgat 5340
tctgagtggt tcagctggct cgagcacacc aagatccgat ggaacggatc atggtaagcg 5400
ctgccactgg ggtgatgaac tccctcctca agaagctcgc ggaactgttg agcgacgagt 5460
acaagctgca gaaatccgtg aagagaaaaa ttcgctcact ggaactcgag ctgagcagca 5520
tcaatgcttt tctcaggaat ttggctgaca aggaggatct ggacccgcag actaaggaat 5580
ggagagacca ggtgagagag atggcctacg aaatcgaaga ctgcatcgac aagtacatgc 5640
acaagctcaa ccacgaaccg aataaagtag gtggcatcaa gggcttcatc agcaagagca 5700
ttgccaaggt gaaaaatatg ggagttgtcc atgggatttc tgatcagctt gagcaactca 5760
agctccaagt cgttgagaca agcgagcgac acaagaggtt actaatgcct gcgcaagtaa 5820
catctggggt atcaaccaca acaattgacc ctcgaatgcc tgcgctgtat gctgatgcta 5880
ctgatcttgt tggtattgat gcaacaagag atgagctcat cgagctggtg accaaccagg 5940
aagagaaaga gttgaaggtg gtgtcgattg ttgggtatgg aggcttggga aagaccacac 6000
ttgcgatcca ggtatacaga caccttcatg ggcaatttga ttttcaagct aaggtgctga 6060
tgtcaagaat ctttgatatg aagaggatac tacgggcaat actctttcaa actaatgaga 6120
cagattacct ggaccagaac acagaatcat ggggtgaaga tttactcatt gagaagctga 6180
gaaaatttct gatggataaa aggtacctaa ttaataatac tctatctagc tggatgaaaa 6240
cttttgctta tagtcaaata tttcccttat catctcacaa ctcatgtgat gaagatcccg 6300
acaacatgtg ccattgattt ttagaaaaca atctcataaa agcacctccg gcaacccctt 6360
gatctctgac aatatgtgcc attgatattg tatacaacat gtaatctatt gtatagagcc 6420
ggtccataca acttgtccct tatttgggtg cagtcaaaca tagtagtgta cctccaaatc 6480
cgacatatat aactaaagct tattttcatt tttgttctac tggatctctg attctgtagg 6540
aaaaataagc tagattaact gaatcctaac cctcgttcat cttcttgctg gaactctaac 6600
cctgtttact cttgtaggta ctttgttgta attgatgaca tatgggatgc tcgaaactgg 6660
gatgccatca aatgcgcttt tcctgatggc aagcgtggaa gtagaataat gacgacaaca 6720
agaatcaata gtgttgctaa gtcatgctgc acccatcgcc gtgatcatat acacaaatta 6780
agtgttctta gtgaagctga ttctcagtgt ttattttata gaagagcttt ttactgtgaa 6840
gatggatgtc cacctgaact ggaagaagtt gccactgaaa ttgtgcaaag atgtggtggt 6900
ctaccattgg cgataattac tctggccagt ttattgagta ctaaatcata tacaagacga 6960
gagtggatga tagtacagga ttctattggt ttgggactta tgaacaatga tgggatggaa 7020
gacatgaata agatattatc tcttagttat attgatcttc cttaccacct gaaaacgtgt 7080
ttgctgtatc ttagtttatt tccagaagac tttctgatta ctagggatcg attagtaaga 7140
aggtggatag cagaagggtt cattacagca gaatgtggga aaactttaga agaacaaggt 7200
gaaagctatt ttaatgagct tataaataga aatttgatcc aaccaataga tatcaagtat 7260
gatggtcgag caagggcatg ccgtgtgcat gatatgattc ttgatttcat tgtatcgaag 7320
gcggctgaag aaaaatttgc aagtttgatt catcaaaagg atgcagttga ctcacgtttt 7380
aaggttcggc gactctcgct taattatggt tcccaaaaag agttgtgcac ggaatcactc 7440
attgtttctc aagctcgatc cctcagtatc tttgggaatt ctgaacagtt gcctcctctt 7500
tcaaacttca gtgcactgag ggtacttgat atagaaagcc atatccagaa cagttatctg 7560
gtgaatattg gagagttgct tcagttgaaa tatatacggc tttctgcaag cactattaca 7620
gagcttccgg aaaacattgg gcaactaaaa tctttggaga caatggatct gaaaaaaact 7680
gatataaaag aactgccagc aagcattgtt caactgcaac gattgaaaca tctactggtt 7740
caaaatgtaa agttgcctgc cggaattgac aagatgcttt ctctccagga cctgtcagaa 7800
ctaattgtag acgatagctg taaagtaact tctttgctag agctgagaac tttggccaat 7860
ttgagttctc ttggccttgt ttggcgcatc agtgattcac acatggaaaa aacaaagttt 7920
gcggacagtt tgttattggc cctctgtgac cttgccaaat ccaaacttca gtttttaaag 7980
gttaccggtg ctgggtctga tgcttcatat gaattcatgt ttgatccttt ctcctcaact 8040
cctcatcgtc tccaagagtt gaccttatac ccaaactgtt atattggtga gaatccaagt 8100
tggatggcct caatggttaa cctcaccaaa ttgaacatta tggttaatcc agtgacacag 8160
gaagctcttg acatttttgg caacttgcct gtcctgctat ttcttgagct gagctcaaaa 8220
gtaatagttc caaaagggct tatcatcgag agtggcacat tcaaatgcct aaaggtgttt 8280
ggcttgtatt gtccggatat tgagagaggg ctgatgttca aagctggagc catgcagagc 8340
attgaaaatt ttacacttcc attcagcgca catgaaccgc aacctatatt gggtgacaat 8400
gattttggca ttccccatct tcgtacgttg cagcaccttg aagttcggat tagctgcaag 8460
ggagcggtgg cttgggaggt ggaaatgttg gctaatgtca ttaggaaggc tgtcattgaa 8520
ctccccagca atccagagcc acaaatcttg aggtattatg tgaaggagat gatacatgat 8580
gacagggaga agagtacaga ggaagctacc ggggactccc tgtatggcac agaatgttct 8640
cacggtatgg aggatcattc ttatggttat tggtatccgc ttccaccgcc tccacgctat 8700
ttcgtccata ttgctgagga ggatccaaat tcttgctcaa tctgctgaag tgtggaatgc 8760
tgcatcaagg tatcatcact cgtttatgca ctttggtaca tttttcagat gctttttggg 8820
tacttgacta gttgctcttt ctgaatagga tagttgaagt atgaaaccaa taatagcctg 8880
ggtcctgcga ctgttcatag agagtctggc ggaccagctc gaggttgtgc ggagcgtcat 8940
tggattgatt tcgttggctg tagttcttta gctagttgtc atttcgctgg ttctgtaata 9000
agttacattc ccgcctggct gttgctggtt ttgtgatgct ggatatataa tgtttcgcct 9060
ccaagtttag atgtaaacta tagctattta tgttaatcct tggaggggag ctgaacctga 9120
ctgaggccgg tggtttgcat tatcagtttt cagaaatgaa cttttagtag ctatgctcct 9180
tataagtcga aaaaagaaca aatacactat tttgtctcct aatgtgctgt ggcccgtttg 9240
ctaatcctac cttattattg ataagcaaaa cacaagatga tggtgatgca ggctagatcg 9300
aggccgccgc cccgagatcc ccatgtcgcc cgaggcgcca gcggtcagat ctgaagcaga 9360
tcgaccacgg ccaccatagg cgtgccccga acggcccccc acgcgaccac cggggaagca 9420
gctcctgccg tcgccccctg agccctcgct gacgcgcccc gtcgccaact accgccccag 9480
cgtcggcggg caggagagag ggccgagggc ttggcggcgc tagggtcccc ccctggtcgc 9540
cgcgcgggga tgactcacgg gagggaaggg gagcagtttg tttagggttg tatctagatt 9600
tcgtaaagtc cttttaaaat gggagagatt ttcacttccc agcctcttta ccaatcaggg 9660
atgcatatgg ccattttagt atgagtacca agataaaatg gtcctcagaa tttttttaaa 9720
tgagtatcaa gatatttctt gatgataggt tgcaccacac accaagcttg atcctcaata 9780
aacaggggaa acccctatgc atcacctatc aattctagga attgaactcg ggtcgctagg 9840
ctgcacaacc gcacgcccaa ccactaagcc aaggctctct ttgcaacgct ccagccttct 9900
agacacggat atacaccaat ggaaaagcaa tgtcatatat agaagaaatt attttgccca 9960
ttgcttttga aggttctagc cttcaatatt gcacagcatg aacacaaaat aggcttcaat 10020
tttgaaagtt tcaggaaacc aaatgcactc aagtcgtaag cgtttatact tgtggatccc 10080
cttggtagcc tatgggagac aagcatacaa aagaccttct ataacagtct gagtctgtac 10140
caagaggccg atgataacta cagacaatat catcgacaca aagtgaaaca caatttgaga 10200
tagaccttaa gagtaagcta ccataagtta gaagattcca aaattaagtc actacaagaa 10260
ttacatttct tcagaaacgt attcagttgg tagagacact aagtggcatt gcatcaaaca 10320
gttggtgggc atatgatcac tgacatgaaa gtatcaccaa agttttgtat gtactagcac 10380
atatgcccgt ccgttgcaat gggagataaa ataatttgca tttaaagtca ctgagaatta 10440
tatgtgcaag taaaatttcg tgtgcacatg aaaaatgaac atttagcttc ttggtttcac 10500
cgtgtgtcaa gaaatcaatc cgctattttc tcattcattg atcgagggca tttaaaagtt 10560
ttgttacgtc attgtgcgcc agagaaggcc catgagttat gcaatctaca cttcctttca 10620
atcctttcaa gcgaggggat ttaatggtat aaagttttta aatattccaa atattctagg 10680
taagatgaac aattttttaa atcctgaaca gtttttaaaa attaatgtat actttcgaaa 10740
aaacgtaaaa aaaatatgaa agggaacatc ttttaaattt gacaaacatt ttttgaaaac 10800
actaaatttt ctataaaaac ataaacattt ttgaatttgt gaacattttt ttacaggaac 10860
atgtgttgaa aattcataag attttaagaa aatgtgtatc ctttatacac gaacattatt 10920
tttaattttg aaaattcact aaaacaagaa tatatttcga atttggagca ttttttaaat 10980
gccatatttt tttacaaatc ttgaacaatt tttaaagaca attatttttt gaaaaaatgg 11040
gaaacttttt taaagttccg aataattttc aaaatagcaa caaaattttg gaattttgaa 11100
catttttttt acttctgaat gttttcgaga atttttaatt taataaataa attctaaaag 11160
aaaaataaac ttgcaaggga aaaaagagat agggaaagga gaataaaaat agatgtaaaa 11220
taccgaaaaa atgaaaaatg ggccagccaa ttattggttg tcctgtgcga acctccgact 11280
attcatcgct cggtgcagaa aatagaattt tccaagatgc atgggcagaa aataagtggg 11340
ccactttgct gggccacaac gcgcgtccca tgtacgtaaa tattttctag cagacaagcg 11400
catgttacta aaaaaaagca gacaaatgca taagaattta gtaccacttt ggatagaaaa 11460
aattcttttc ggtgaacaga tgaaaaaatt ggcgaaacgc accttgcttt attagtaggt 11520
atagaaacgg gccatcggca tggtcagaga tggagaaggc ggagtcagcg tccacccagt 11580
tcaggggagg gatatgtgga gggggcgttg gagggagagg tgcatatata ggctgcgccg 11640
cgtgagggac gcttcaggaa aggatctgca tgacaggggc gctaggtata gccaaacaag 11700
gtacgcaccg ggaaaggtgc cgatagaaag 11730
<210> 2
<211> 2955
<212> DNA
<213> Artificial sequence
<400> 2
atggaacgga tcatggtaag cgctgccact ggggtgatga actccctcct caagaagctc 60
gcggaactgt tgagcgacga gtacaagctg cagaaatccg tgaagagaaa aattcgctca 120
ctggaactcg agctgagcag catcaatgct tttctcagga atttggctga caaggaggat 180
ctggacccgc agactaagga atggagagac caggtgagag agatggccta cgaaatcgaa 240
gactgcatcg acaagtacat gcacaagctc aaccacgaac cgaataaagt aggtggcatc 300
aagggcttca tcagcaagag cattgccaag gtgaaaaata tgggagttgt ccatgggatt 360
tctgatcagc ttgagcaact caagctccaa gtcgttgaga caagcgagcg acacaagagg 420
ttactaatgc ctgcgcaagt aacatctggg gtatcaacca caacaattga ccctcgaatg 480
cctgcgctgt atgctgatgc tactgatctt gttggtattg atgcaacaag agatgagctc 540
atcgagctgg tgaccaacca ggaagagaaa gagttgaagg tggtgtcgat tgttgggtat 600
ggaggcttgg gaaagaccac acttgcgatc caggtataca gacaccttca tgggcaattt 660
gattttcaag ctaaggtgct gatgtcaaga atctttgata tgaagaggat actacgggca 720
atactctttc aaactaatga gacagattac ctggaccaga acacagaatc atggggtgaa 780
gatttactca ttgagaagct gagaaaattt ctgatggata aaaggtactt tgttgtaatt 840
gatgacatat gggatgctcg aaactgggat gccatcaaat gcgcttttcc tgatggcaag 900
cgtggaagta gaataatgac gacaacaaga atcaatagtg ttgctaagtc atgctgcacc 960
catcgccgtg atcatataca caaattaagt gttcttagtg aagctgattc tcagtgttta 1020
ttttatagaa gagcttttta ctgtgaagat ggatgtccac ctgaactgga agaagttgcc 1080
actgaaattg tgcaaagatg tggtggtcta ccattggcga taattactct ggccagttta 1140
ttgagtacta aatcatatac aagacgagag tggatgatag tacaggattc tattggtttg 1200
ggacttatga acaatgatgg gatggaagac atgaataaga tattatctct tagttatatt 1260
gatcttcctt accacctgaa aacgtgtttg ctgtatctta gtttatttcc agaagacttt 1320
ctgattacta gggatcgatt agtaagaagg tggatagcag aagggttcat tacagcagaa 1380
tgtgggaaaa ctttagaaga acaaggtgaa agctatttta atgagcttat aaatagaaat 1440
ttgatccaac caatagatat caagtatgat ggtcgagcaa gggcatgccg tgtgcatgat 1500
atgattcttg atttcattgt atcgaaggcg gctgaagaaa aatttgcaag tttgattcat 1560
caaaaggatg cagttgactc acgttttaag gttcggcgac tctcgcttaa ttatggttcc 1620
caaaaagagt tgtgcacgga atcactcatt gtttctcaag ctcgatccct cagtatcttt 1680
gggaattctg aacagttgcc tcctctttca aacttcagtg cactgagggt acttgatata 1740
gaaagccata tccagaacag ttatctggtg aatattggag agttgcttca gttgaaatat 1800
atacggcttt ctgcaagcac tattacagag cttccggaaa acattgggca actaaaatct 1860
ttggagacaa tggatctgaa aaaaactgat ataaaagaac tgccagcaag cattgttcaa 1920
ctgcaacgat tgaaacatct actggttcaa aatgtaaagt tgcctgccgg aattgacaag 1980
atgctttctc tccaggacct gtcagaacta attgtagacg atagctgtaa agtaacttct 2040
ttgctagagc tgagaacttt ggccaatttg agttctcttg gccttgtttg gcgcatcagt 2100
gattcacaca tggaaaaaac aaagtttgcg gacagtttgt tattggccct ctgtgacctt 2160
gccaaatcca aacttcagtt tttaaaggtt accggtgctg ggtctgatgc ttcatatgaa 2220
ttcatgtttg atcctttctc ctcaactcct catcgtctcc aagagttgac cttataccca 2280
aactgttata ttggtgagaa tccaagttgg atggcctcaa tggttaacct caccaaattg 2340
aacattatgg ttaatccagt gacacaggaa gctcttgaca tttttggcaa cttgcctgtc 2400
ctgctatttc ttgagctgag ctcaaaagta atagttccaa aagggcttat catcgagagt 2460
ggcacattca aatgcctaaa ggtgtttggc ttgtattgtc cggatattga gagagggctg 2520
atgttcaaag ctggagccat gcagagcatt gaaaatttta cacttccatt cagcgcacat 2580
gaaccgcaac ctatattggg tgacaatgat tttggcattc cccatcttcg tacgttgcag 2640
caccttgaag ttcggattag ctgcaaggga gcggtggctt gggaggtgga aatgttggct 2700
aatgtcatta ggaaggctgt cattgaactc cccagcaatc cagagccaca aatcttgagg 2760
tattatgtga aggagatgat acatgatgac agggagaaga gtacagagga agctaccggg 2820
gactccctgt atggcacaga atgttctcac ggtatggagg atcattctta tggttattgg 2880
tatccgcttc caccgcctcc acgctatttc gtccatattg ctgaggagga tccaaattct 2940
tgctcaatct gctga 2955
<210> 3
<211> 984
<212> PRT
<213> Artificial sequence
<400> 3
Met Glu Arg Ile Met Val Ser Ala Ala Thr Gly Val Met Asn Ser Leu
1 5 10 15
Leu Lys Lys Leu Ala Glu Leu Leu Ser Asp Glu Tyr Lys Leu Gln Lys
20 25 30
Ser Val Lys Arg Lys Ile Arg Ser Leu Glu Leu Glu Leu Ser Ser Ile
35 40 45
Asn Ala Phe Leu Arg Asn Leu Ala Asp Lys Glu Asp Leu Asp Pro Gln
50 55 60
Thr Lys Glu Trp Arg Asp Gln Val Arg Glu Met Ala Tyr Glu Ile Glu
65 70 75 80
Asp Cys Ile Asp Lys Tyr Met His Lys Leu Asn His Glu Pro Asn Lys
85 90 95
Val Gly Gly Ile Lys Gly Phe Ile Ser Lys Ser Ile Ala Lys Val Lys
100 105 110
Asn Met Gly Val Val His Gly Ile Ser Asp Gln Leu Glu Gln Leu Lys
115 120 125
Leu Gln Val Val Glu Thr Ser Glu Arg His Lys Arg Leu Leu Met Pro
130 135 140
Ala Gln Val Thr Ser Gly Val Ser Thr Thr Thr Ile Asp Pro Arg Met
145 150 155 160
Pro Ala Leu Tyr Ala Asp Ala Thr Asp Leu Val Gly Ile Asp Ala Thr
165 170 175
Arg Asp Glu Leu Ile Glu Leu Val Thr Asn Gln Glu Glu Lys Glu Leu
180 185 190
Lys Val Val Ser Ile Val Gly Tyr Gly Gly Leu Gly Lys Thr Thr Leu
195 200 205
Ala Ile Gln Val Tyr Arg His Leu His Gly Gln Phe Asp Phe Gln Ala
210 215 220
Lys Val Leu Met Ser Arg Ile Phe Asp Met Lys Arg Ile Leu Arg Ala
225 230 235 240
Ile Leu Phe Gln Thr Asn Glu Thr Asp Tyr Leu Asp Gln Asn Thr Glu
245 250 255
Ser Trp Gly Glu Asp Leu Leu Ile Glu Lys Leu Arg Lys Phe Leu Met
260 265 270
Asp Lys Arg Tyr Phe Val Val Ile Asp Asp Ile Trp Asp Ala Arg Asn
275 280 285
Trp Asp Ala Ile Lys Cys Ala Phe Pro Asp Gly Lys Arg Gly Ser Arg
290 295 300
Ile Met Thr Thr Thr Arg Ile Asn Ser Val Ala Lys Ser Cys Cys Thr
305 310 315 320
His Arg Arg Asp His Ile His Lys Leu Ser Val Leu Ser Glu Ala Asp
325 330 335
Ser Gln Cys Leu Phe Tyr Arg Arg Ala Phe Tyr Cys Glu Asp Gly Cys
340 345 350
Pro Pro Glu Leu Glu Glu Val Ala Thr Glu Ile Val Gln Arg Cys Gly
355 360 365
Gly Leu Pro Leu Ala Ile Ile Thr Leu Ala Ser Leu Leu Ser Thr Lys
370 375 380
Ser Tyr Thr Arg Arg Glu Trp Met Ile Val Gln Asp Ser Ile Gly Leu
385 390 395 400
Gly Leu Met Asn Asn Asp Gly Met Glu Asp Met Asn Lys Ile Leu Ser
405 410 415
Leu Ser Tyr Ile Asp Leu Pro Tyr His Leu Lys Thr Cys Leu Leu Tyr
420 425 430
Leu Ser Leu Phe Pro Glu Asp Phe Leu Ile Thr Arg Asp Arg Leu Val
435 440 445
Arg Arg Trp Ile Ala Glu Gly Phe Ile Thr Ala Glu Cys Gly Lys Thr
450 455 460
Leu Glu Glu Gln Gly Glu Ser Tyr Phe Asn Glu Leu Ile Asn Arg Asn
465 470 475 480
Leu Ile Gln Pro Ile Asp Ile Lys Tyr Asp Gly Arg Ala Arg Ala Cys
485 490 495
Arg Val His Asp Met Ile Leu Asp Phe Ile Val Ser Lys Ala Ala Glu
500 505 510
Glu Lys Phe Ala Ser Leu Ile His Gln Lys Asp Ala Val Asp Ser Arg
515 520 525
Phe Lys Val Arg Arg Leu Ser Leu Asn Tyr Gly Ser Gln Lys Glu Leu
530 535 540
Cys Thr Glu Ser Leu Ile Val Ser Gln Ala Arg Ser Leu Ser Ile Phe
545 550 555 560
Gly Asn Ser Glu Gln Leu Pro Pro Leu Ser Asn Phe Ser Ala Leu Arg
565 570 575
Val Leu Asp Ile Glu Ser His Ile Gln Asn Ser Tyr Leu Val Asn Ile
580 585 590
Gly Glu Leu Leu Gln Leu Lys Tyr Ile Arg Leu Ser Ala Ser Thr Ile
595 600 605
Thr Glu Leu Pro Glu Asn Ile Gly Gln Leu Lys Ser Leu Glu Thr Met
610 615 620
Asp Leu Lys Lys Thr Asp Ile Lys Glu Leu Pro Ala Ser Ile Val Gln
625 630 635 640
Leu Gln Arg Leu Lys His Leu Leu Val Gln Asn Val Lys Leu Pro Ala
645 650 655
Gly Ile Asp Lys Met Leu Ser Leu Gln Asp Leu Ser Glu Leu Ile Val
660 665 670
Asp Asp Ser Cys Lys Val Thr Ser Leu Leu Glu Leu Arg Thr Leu Ala
675 680 685
Asn Leu Ser Ser Leu Gly Leu Val Trp Arg Ile Ser Asp Ser His Met
690 695 700
Glu Lys Thr Lys Phe Ala Asp Ser Leu Leu Leu Ala Leu Cys Asp Leu
705 710 715 720
Ala Lys Ser Lys Leu Gln Phe Leu Lys Val Thr Gly Ala Gly Ser Asp
725 730 735
Ala Ser Tyr Glu Phe Met Phe Asp Pro Phe Ser Ser Thr Pro His Arg
740 745 750
Leu Gln Glu Leu Thr Leu Tyr Pro Asn Cys Tyr Ile Gly Glu Asn Pro
755 760 765
Ser Trp Met Ala Ser Met Val Asn Leu Thr Lys Leu Asn Ile Met Val
770 775 780
Asn Pro Val Thr Gln Glu Ala Leu Asp Ile Phe Gly Asn Leu Pro Val
785 790 795 800
Leu Leu Phe Leu Glu Leu Ser Ser Lys Val Ile Val Pro Lys Gly Leu
805 810 815
Ile Ile Glu Ser Gly Thr Phe Lys Cys Leu Lys Val Phe Gly Leu Tyr
820 825 830
Cys Pro Asp Ile Glu Arg Gly Leu Met Phe Lys Ala Gly Ala Met Gln
835 840 845
Ser Ile Glu Asn Phe Thr Leu Pro Phe Ser Ala His Glu Pro Gln Pro
850 855 860
Ile Leu Gly Asp Asn Asp Phe Gly Ile Pro His Leu Arg Thr Leu Gln
865 870 875 880
His Leu Glu Val Arg Ile Ser Cys Lys Gly Ala Val Ala Trp Glu Val
885 890 895
Glu Met Leu Ala Asn Val Ile Arg Lys Ala Val Ile Glu Leu Pro Ser
900 905 910
Asn Pro Glu Pro Gln Ile Leu Arg Tyr Tyr Val Lys Glu Met Ile His
915 920 925
Asp Asp Arg Glu Lys Ser Thr Glu Glu Ala Thr Gly Asp Ser Leu Tyr
930 935 940
Gly Thr Glu Cys Ser His Gly Met Glu Asp His Ser Tyr Gly Tyr Trp
945 950 955 960
Tyr Pro Leu Pro Pro Pro Pro Arg Tyr Phe Val His Ile Ala Glu Glu
965 970 975
Asp Pro Asn Ser Cys Ser Ile Cys
980
<210> 4
<211> 214
<212> DNA
<213> Artificial sequence
<400> 4
cgggtacatc tgactgttca atttacagtc ttctttctca gatctgaacc tactctgctc 60
ttggtagcag cagcttgtac ttcttctcaa gtcaccacgg cagtatctac actcttagat 120
tctgtgattc tgagtggttc agctggctcg agcacaccaa gatccgatgg aacggatcat 180
ggtaagcgct gccactgggg tgatgaactc cctc 214
<210> 5
<211> 214
<212> DNA
<213> Artificial sequence
<400> 5
cgggtacatc tgactgttca atttacagtc ttctttctca gatctgaacc tactctgctc 60
ttggtagcag cagcttgtac ttcttctcaa gtcaccacgg cagtatctac actcttagat 120
tctgtgattc tgagtggttc agctggctcg agcacaccaa gatccgatgg aacggatcat 180
ggtgagcact gccactgggg tgatgaactc cctc 214
<210> 6
<211> 214
<212> DNA
<213> Artificial sequence
<400> 6
cgggtacatc tgactgttca atttacagtc ttctttctca gatctgaacc tactctgctc 60
ttggtagcag cagcttgtac ttcttctcaa gtcaccacgg cagtatctac actcttagat 120
tctgtgattc tgagtggttc agctggctcg agcacaccaa gatccgatgg aacggatcat 180
ggtaagcttt gccactgggg tgatgaactc cctc 214
<210> 7
<211> 214
<212> DNA
<213> Artificial sequence
<400> 7
cgggtacatc tgactgttca atttacagtc ttctttctca gatctgaacc tactctgctc 60
ttggtagcag cagcttgtac ttcttctcaa gtcaccacgg cagtatctac actcttagat 120
tctgtgattc tgagtggttc agctggctcg agcacaccaa gatccgatgg aacggatcat 180
ggtgagcttt gccactgggg tgatgaactc cctc 214
<210> 8
<211> 27
<212> DNA
<213> Artificial sequence
<400> 8
cgggtacatc tgactgttca atttaca 27
<210> 9
<211> 30
<212> DNA
<213> Artificial sequence
<400> 9
gagggagttc atcaccccag tggcaaagct 30
Claims (7)
1.特异引物对的应用，为如下b1）-b3）中的任一种：
b1）辅助鉴定待测小麦是否具有白粉病抗性；
b2）辅助筛选抗白粉病的小麦品种；
b3）辅助筛选感白粉病的小麦品种；
所述特异引物对由引物F和引物R组成；
所述引物F为序列表的序列8所示的单链DNA分子；
所述引物R为序列表的序列9所示的单链DNA分子。
2.DNA片段的应用，为如下b1）-b3）中的任一种：
b1）辅助鉴定待测小麦是否具有白粉病抗性；
b2）辅助筛选抗白粉病的小麦品种；
b3）辅助筛选感白粉病的小麦品种；
所述DNA片段的核苷酸序列如序列表中序列6或序列表中序列7所示。
3.根据权利要求1或2所述的应用，其特征在于：所述白粉病为白粉菌引起的病害。
4.一种检测待测小麦是否具有白粉病抗性的方法，为方法R1）或方法R2）：
方法R1）检测待测小麦的基因组中是否含有序列表中序列4所示的DNA片段和序列表中的序列5所示的DNA片段，如果待测小麦的基因组中含有序列表中序列4所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；
方法R2）检测待测小麦的基因组中是否含有序列表中序列4所示的DNA片段和序列表中的序列5所示的DNA片段，然后进行如下判断：如果待测小麦的基因组中含有序列表中序列4所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果待测小麦的基因组中含有序列表中序列5所示的DNA片段且不含有序列表中序列4所示的DNA片段，则待测小麦不具有或疑似不具有白粉病抗性。
5.一种检测待测小麦是否具有白粉病抗性的方法，为方法S1）或方法S2）：
方法S1）以待测小麦的基因组DNA为模板，采用权利要求1中所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，如果酶切产物中具有189bp的DNA片段，则待测小麦具有或疑似具有白粉病抗性；
方法S2）以待测小麦的基因组DNA为模板，采用权利要求1中所述特异引物对进行PCR扩增，得到PCR扩增产物；用限制性内切酶HindⅢ酶切所述PCR扩增产物，然后进行如下判断：如果酶切产物中具有189bp的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果酶切产物中“具有214bp的DNA片段且不具有189bp的DNA片段”或“不具有任何DNA片段”，则待测小麦不具有或疑似不具有白粉病抗性。
6.一种检测待测小麦是否具有白粉病抗性的方法，为方法T1）或方法T2）：
方法T1）以待测小麦的基因组DNA为模板，采用权利要求1中所述特异引物对进行PCR扩增，得到PCR扩增产物；将所述PCR扩增产物进行测序，然后进行如下判断：如果PCR扩增产物含有序列表中序列6所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；
方法T2）以待测小麦的基因组DNA为模板，采用权利要求1中所述特异引物对进行PCR扩增，得到PCR扩增产物；将所述PCR扩增产物进行测序，然后进行如下判断：如果PCR扩增产物含有序列表中序列6所示的DNA片段，则待测小麦具有或疑似具有白粉病抗性；如果PCR扩增产物“含有序列表中序列7所示的DNA片段且不含有序列表中序列6所示的DNA片段”或“不含有任何DNA片段”，则待测小麦不具有或疑似不具有白粉病抗性。
7.根据权利要求4至6任一所述的方法，其特征在于：所述白粉病为白粉菌引起的病害。
Priority Applications (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN201910602364.XA CN112176083B (zh) | 2019-07-05 | 2019-07-05 | 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 |
Applications Claiming Priority (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN201910602364.XA CN112176083B (zh) | 2019-07-05 | 2019-07-05 | 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 |
Publications (2)
| Publication Number | Publication Date |
|---|---|
| CN112176083A CN112176083A (zh) | 2021-01-05 |
| CN112176083B true CN112176083B (zh) | 2022-04-12 |
Family
ID=73914595
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| CN201910602364.XA Active CN112176083B (zh) | 2019-07-05 | 2019-07-05 | 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 |
Country Status (1)
| Country | Link |
|---|---|
| CN (1) | CN112176083B (zh) |
Families Citing this family (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN111732644B (zh) * | 2019-03-20 | 2021-11-16 | 中国科学院遗传与发育生物学研究所 | 抗白粉病相关蛋白Pm41及其编码基因与应用 |
| CN113652423B (zh) * | 2021-09-29 | 2023-06-02 | 西北农林科技大学 | 一种辅助鉴定待测小麦抗旱性的方法及其专用的分子标记 |
Citations (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN1428440A (zh) * | 2002-09-27 | 2003-07-09 | 天津师范大学 | 与小麦抗白粉病基因相连锁的分子标记 |
| CN102533750A (zh) * | 2012-02-20 | 2012-07-04 | 中国科学院成都生物研究所 | 一种小麦抗白粉病基因分子标记引物及其用途 |
| CN108950057A (zh) * | 2018-08-30 | 2018-12-07 | 中国科学院遗传与发育生物学研究所 | 乌拉尔图小麦抗白粉病基因Pm60特异分子标记的开发及其应用 |
-
2019
- 2019-07-05 CN CN201910602364.XA patent/CN112176083B/zh active Active
Patent Citations (3)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN1428440A (zh) * | 2002-09-27 | 2003-07-09 | 天津师范大学 | 与小麦抗白粉病基因相连锁的分子标记 |
| CN102533750A (zh) * | 2012-02-20 | 2012-07-04 | 中国科学院成都生物研究所 | 一种小麦抗白粉病基因分子标记引物及其用途 |
| CN108950057A (zh) * | 2018-08-30 | 2018-12-07 | 中国科学院遗传与发育生物学研究所 | 乌拉尔图小麦抗白粉病基因Pm60特异分子标记的开发及其应用 |
Non-Patent Citations (1)
| Title |
|---|
| Triticum turgidum subsp. durum genome assembly, chromosome: chr3B;LT934116.1;《GenBank 》;20190211;全文 * |
Also Published As
| Publication number | Publication date |
|---|---|
| CN112176083A (zh) | 2021-01-05 |
Similar Documents
| Publication | Publication Date | Title |
|---|---|---|
| Xu et al. | High-resolution mapping of loci conferring resistance to sugarcane mosaic virus in maize using RFLP, SSR, and AFLP markers | |
| CN111719008B (zh) | 一种与小麦白粉病抗病基因Pm5e共分离的SNP及其应用 | |
| CN112176083B (zh) | 小麦抗白粉病相关基因Pm41的功能型分子标记及其应用 | |
| Yang et al. | The genome of the rice variety LTH provides insight into its universal susceptibility mechanism to worldwide rice blast fungal strains | |
| CN112813192B (zh) | 小麦抗叶锈病基因Lr13的分子标记HBAU-LrZH22及其检测引物和应用 | |
| CN104404054A (zh) | 玉米抗丝黑穗病相关基因ZmNL的克隆鉴定及其在抗丝黑穗病玉米育种中的用途 | |
| CN113234733B (zh) | 一个水稻叶绿体发育的相关基因及分子标记和应用 | |
| Azhaguvel et al. | Methodological advancement in molecular markers to delimit the gene (s) for crop improvement | |
| Oh et al. | Fine mapping in tomato using microsynteny with the Arabidopsis genome: the Diageotropica (Dgt) locus | |
| Nagy et al. | Fine mapping of the Pc locus of Sorghum bicolor, a gene controlling the reaction to a fungal pathogen and its host-selective toxin | |
| CN111732644B (zh) | 抗白粉病相关蛋白Pm41及其编码基因与应用 | |
| KR101410329B1 (ko) | Ｔｓｗｖ 저항성 고추 품종을 선별하기 위한 프라이머 세트, 방법 및 키트 | |
| CN112760319B (zh) | 高大山羊草2Sl染色体特异分子标记的开发及其应用 | |
| KR102429219B1 (ko) | 콩 역병균에 대한 저항성 또는 감수성을 가지는 콩 품종 판별용 마커 조성물 및 이의 용도 | |
| Babu et al. | Isolation, fine mapping and expression profiling of a lesion mimic genotype, spl NF4050-8 that confers blast resistance in rice | |
| CN114350687A (zh) | 一种水稻抗白叶枯病基因、蛋白及其应用 | |
| CN114480705A (zh) | 一种水稻抗白叶枯病基因xa23的snp分子标记及其扩增引物和应用 | |
| CN107653341B (zh) | 一种检测粗山羊草抗白粉病基因的kasp标记及应用 | |
| KR101432281B1 (ko) | 캡시컴 안늄의 종내집단을 이용한 ｓｓｒ 마커 및 유전자 연관지도 | |
| KR101401980B1 (ko) | 역병 저항성 고추 품종을 선별하기 위한 특이 프라이머 및 이의 용도 | |
| Sugihara et al. | Tangled gene-for-gene interactions mediate co-evolution of the rice NLR immune receptor Pik and blast fungus effector proteins | |
| CN108728563B (zh) | 大白菜Bra013400移码突变的InDel标记及其在育种实践中的应用 | |
| CN106636386B (zh) | 与玉米抗丝黑穗病次主效位点连锁的分子标记DNdCAPS8.03-1及其应用 | |
| KR100984169B1 (ko) | Ｔｍｖ 저항성 고추 품종을 선별하기 위한 프라이머 세트,방법 및 키트 | |
| Mráček | Investigation of interspecific genome-plastome incompatibility in Oenothera and Passiflora |
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 | ||
| GR01 | Patent grant | ||
| GR01 | Patent grant |