CN114106120B - 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 - Google Patents
一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 Download PDFInfo
- Publication number
- CN114106120B CN114106120B CN202111154498.3A CN202111154498A CN114106120B CN 114106120 B CN114106120 B CN 114106120B CN 202111154498 A CN202111154498 A CN 202111154498A CN 114106120 B CN114106120 B CN 114106120B
- Authority
- CN
- China
- Prior art keywords
- plant
- phhma5ii
- protein
- heavy metal
- petunia
- 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
- 229910001385 heavy metal Inorganic materials 0.000 title claims abstract description 57
- 108010078791 Carrier Proteins Proteins 0.000 title claims abstract description 13
- 102000014914 Carrier Proteins Human genes 0.000 title claims abstract description 9
- 108090000623 proteins and genes Proteins 0.000 claims abstract description 96
- 241000207748 Petunia Species 0.000 claims abstract description 75
- 102000004169 proteins and genes Human genes 0.000 claims abstract description 46
- XEEYBQQBJWHFJM-UHFFFAOYSA-N Iron Chemical compound [Fe] XEEYBQQBJWHFJM-UHFFFAOYSA-N 0.000 claims abstract description 37
- RYGMFSIKBFXOCR-UHFFFAOYSA-N Copper Chemical compound [Cu] RYGMFSIKBFXOCR-UHFFFAOYSA-N 0.000 claims abstract description 35
- 229910052802 copper Inorganic materials 0.000 claims abstract description 35
- 239000010949 copper Substances 0.000 claims abstract description 35
- 229910052742 iron Inorganic materials 0.000 claims abstract description 20
- 108091033409 CRISPR Proteins 0.000 claims abstract description 18
- 125000003275 alpha amino acid group Chemical group 0.000 claims abstract description 15
- PWHULOQIROXLJO-UHFFFAOYSA-N Manganese Chemical compound [Mn] PWHULOQIROXLJO-UHFFFAOYSA-N 0.000 claims abstract description 10
- 229910052748 manganese Inorganic materials 0.000 claims abstract description 10
- 239000011572 manganese Substances 0.000 claims abstract description 10
- 241000196324 Embryophyta Species 0.000 claims description 89
- 238000000034 method Methods 0.000 claims description 35
- 108091033319 polynucleotide Proteins 0.000 claims description 27
- 102000040430 polynucleotide Human genes 0.000 claims description 27
- 239000002157 polynucleotide Substances 0.000 claims description 27
- 239000013604 expression vector Substances 0.000 claims description 23
- 150000007523 nucleic acids Chemical class 0.000 claims description 18
- 102000039446 nucleic acids Human genes 0.000 claims description 17
- 108020004707 nucleic acids Proteins 0.000 claims description 17
- 238000003259 recombinant expression Methods 0.000 claims description 16
- 230000002452 interceptive effect Effects 0.000 claims description 14
- 230000001737 promoting effect Effects 0.000 claims description 11
- 241000894006 Bacteria Species 0.000 claims description 10
- 230000008635 plant growth Effects 0.000 claims description 10
- 239000013598 vector Substances 0.000 claims description 9
- 239000012620 biological material Substances 0.000 claims description 8
- 108091026890 Coding region Proteins 0.000 claims description 6
- 238000010354 CRISPR gene editing Methods 0.000 claims description 5
- HCHKCACWOHOZIP-UHFFFAOYSA-N Zinc Chemical compound [Zn] HCHKCACWOHOZIP-UHFFFAOYSA-N 0.000 claims description 5
- 239000012634 fragment Substances 0.000 claims description 5
- 239000011701 zinc Substances 0.000 claims description 5
- 229910052725 zinc Inorganic materials 0.000 claims description 5
- 230000003247 decreasing effect Effects 0.000 claims description 4
- 239000002773 nucleotide Substances 0.000 claims description 3
- 125000003729 nucleotide group Chemical group 0.000 claims description 3
- 230000000415 inactivating effect Effects 0.000 claims 3
- 230000002829 reductive effect Effects 0.000 abstract description 7
- 230000012010 growth Effects 0.000 abstract description 5
- 238000010362 genome editing Methods 0.000 abstract description 4
- 125000000539 amino acid group Chemical group 0.000 abstract description 3
- 238000005516 engineering process Methods 0.000 abstract description 3
- 108020004414 DNA Proteins 0.000 description 26
- 241000589158 Agrobacterium Species 0.000 description 19
- 239000000047 product Substances 0.000 description 14
- 239000013612 plasmid Substances 0.000 description 13
- 210000004027 cell Anatomy 0.000 description 12
- 229910000365 copper sulfate Inorganic materials 0.000 description 11
- ARUVKPQLZAKDPS-UHFFFAOYSA-L copper(II) sulfate Chemical group [Cu+2].[O-][S+2]([O-])([O-])[O-] ARUVKPQLZAKDPS-UHFFFAOYSA-L 0.000 description 11
- XLYOFNOQVPJJNP-UHFFFAOYSA-N water Substances O XLYOFNOQVPJJNP-UHFFFAOYSA-N 0.000 description 11
- OJOBTAOGJIWAGB-UHFFFAOYSA-N acetosyringone Chemical compound COC1=CC(C(C)=O)=CC(OC)=C1O OJOBTAOGJIWAGB-UHFFFAOYSA-N 0.000 description 10
- 230000006870 function Effects 0.000 description 9
- 230000009466 transformation Effects 0.000 description 9
- 230000009261 transgenic effect Effects 0.000 description 9
- 238000012408 PCR amplification Methods 0.000 description 8
- 241000219194 Arabidopsis Species 0.000 description 7
- 239000002299 complementary DNA Substances 0.000 description 7
- 239000002609 medium Substances 0.000 description 7
- 239000000203 mixture Substances 0.000 description 7
- 239000003112 inhibitor Substances 0.000 description 6
- 239000007787 solid Substances 0.000 description 6
- 239000000243 solution Substances 0.000 description 6
- 108091032973 (ribonucleotides)n+m Proteins 0.000 description 5
- 239000000463 material Substances 0.000 description 5
- 238000012216 screening Methods 0.000 description 5
- 229920001817 Agar Polymers 0.000 description 4
- 108010077544 Chromatin Proteins 0.000 description 4
- LFQSCWFLJHTTHZ-UHFFFAOYSA-N Ethanol Chemical compound CCO LFQSCWFLJHTTHZ-UHFFFAOYSA-N 0.000 description 4
- 101150078264 HMA5 gene Proteins 0.000 description 4
- 240000004808 Saccharomyces cerevisiae Species 0.000 description 4
- FAPWRFPIFSIZLT-UHFFFAOYSA-M Sodium chloride Chemical compound [Na+].[Cl-] FAPWRFPIFSIZLT-UHFFFAOYSA-M 0.000 description 4
- 239000008272 agar Substances 0.000 description 4
- 238000000137 annealing Methods 0.000 description 4
- 210000003483 chromatin Anatomy 0.000 description 4
- OVBPIULPVIDEAO-LBPRGKRZSA-N folic acid Chemical compound C=1N=C2NC(N)=NC(=O)C2=NC=1CNC1=CC=C(C(=O)N[C@@H](CCC(O)=O)C(O)=O)C=C1 OVBPIULPVIDEAO-LBPRGKRZSA-N 0.000 description 4
- 238000009472 formulation Methods 0.000 description 4
- 230000001404 mediated effect Effects 0.000 description 4
- 239000000843 powder Substances 0.000 description 4
- 230000002786 root growth Effects 0.000 description 4
- 230000001954 sterilising effect Effects 0.000 description 4
- 210000001519 tissue Anatomy 0.000 description 4
- 241000219195 Arabidopsis thaliana Species 0.000 description 3
- 102000016928 DNA-directed DNA polymerase Human genes 0.000 description 3
- 108010014303 DNA-directed DNA polymerase Proteins 0.000 description 3
- WQZGKKKJIJFFOK-GASJEMHNSA-N Glucose Natural products OC[C@H]1OC(O)[C@H](O)[C@@H](O)[C@@H]1O WQZGKKKJIJFFOK-GASJEMHNSA-N 0.000 description 3
- PEDCQBHIVMGVHV-UHFFFAOYSA-N Glycerine Chemical compound OCC(O)CO PEDCQBHIVMGVHV-UHFFFAOYSA-N 0.000 description 3
- KFZMGEQAYNKOFK-UHFFFAOYSA-N Isopropanol Chemical compound CC(C)O KFZMGEQAYNKOFK-UHFFFAOYSA-N 0.000 description 3
- 241000880493 Leptailurus serval Species 0.000 description 3
- 108091028043 Nucleic acid sequence Proteins 0.000 description 3
- 108020004511 Recombinant DNA Proteins 0.000 description 3
- 229930006000 Sucrose Natural products 0.000 description 3
- CZMRCDWAGMRECN-UGDNZRGBSA-N Sucrose Chemical compound O[C@H]1[C@H](O)[C@@H](CO)O[C@@]1(CO)O[C@@H]1[C@H](O)[C@@H](O)[C@H](O)[C@@H](CO)O1 CZMRCDWAGMRECN-UGDNZRGBSA-N 0.000 description 3
- 239000007864 aqueous solution Substances 0.000 description 3
- 230000001580 bacterial effect Effects 0.000 description 3
- 238000006243 chemical reaction Methods 0.000 description 3
- 238000010367 cloning Methods 0.000 description 3
- 238000007796 conventional method Methods 0.000 description 3
- 238000012258 culturing Methods 0.000 description 3
- 238000004925 denaturation Methods 0.000 description 3
- 230000036425 denaturation Effects 0.000 description 3
- 238000001784 detoxification Methods 0.000 description 3
- 230000000694 effects Effects 0.000 description 3
- 239000008103 glucose Substances 0.000 description 3
- 108010050848 glycylleucine Proteins 0.000 description 3
- -1 iron ion Chemical class 0.000 description 3
- 229930027917 kanamycin Natural products 0.000 description 3
- SBUJHOSQTJFQJX-NOAMYHISSA-N kanamycin Chemical compound O[C@@H]1[C@@H](O)[C@H](O)[C@@H](CN)O[C@@H]1O[C@H]1[C@H](O)[C@@H](O[C@@H]2[C@@H]([C@@H](N)[C@H](O)[C@@H](CO)O2)O)[C@H](N)C[C@@H]1N SBUJHOSQTJFQJX-NOAMYHISSA-N 0.000 description 3
- 229960000318 kanamycin Drugs 0.000 description 3
- 229930182823 kanamycin A Natural products 0.000 description 3
- 239000007788 liquid Substances 0.000 description 3
- 230000035772 mutation Effects 0.000 description 3
- 235000015097 nutrients Nutrition 0.000 description 3
- 238000012257 pre-denaturation Methods 0.000 description 3
- 238000012163 sequencing technique Methods 0.000 description 3
- 239000002689 soil Substances 0.000 description 3
- 239000005720 sucrose Substances 0.000 description 3
- 239000006228 supernatant Substances 0.000 description 3
- 108010061238 threonyl-glycine Proteins 0.000 description 3
- 230000001131 transforming effect Effects 0.000 description 3
- SUMYEVXWCAYLLJ-GUBZILKMSA-N Ala-Leu-Gln Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O SUMYEVXWCAYLLJ-GUBZILKMSA-N 0.000 description 2
- HEDRZPFGACZZDS-UHFFFAOYSA-N Chloroform Chemical compound ClC(Cl)Cl HEDRZPFGACZZDS-UHFFFAOYSA-N 0.000 description 2
- 108091033380 Coding strand Proteins 0.000 description 2
- 229920000742 Cotton Polymers 0.000 description 2
- JPVYNHNXODAKFH-UHFFFAOYSA-N Cu2+ Chemical compound [Cu+2] JPVYNHNXODAKFH-UHFFFAOYSA-N 0.000 description 2
- CGOHAEBMDSEKFB-FXQIFTODSA-N Glu-Glu-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O CGOHAEBMDSEKFB-FXQIFTODSA-N 0.000 description 2
- WVYJNPCWJYBHJG-YVNDNENWSA-N Glu-Ile-Gln Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(N)=O)C(O)=O WVYJNPCWJYBHJG-YVNDNENWSA-N 0.000 description 2
- 241000219146 Gossypium Species 0.000 description 2
- MHAJPDPJQMAIIY-UHFFFAOYSA-N Hydrogen peroxide Chemical compound OO MHAJPDPJQMAIIY-UHFFFAOYSA-N 0.000 description 2
- 206010020649 Hyperkeratosis Diseases 0.000 description 2
- RCFDOSNHHZGBOY-UHFFFAOYSA-N L-isoleucyl-L-alanine Natural products CCC(C)C(N)C(=O)NC(C)C(O)=O RCFDOSNHHZGBOY-UHFFFAOYSA-N 0.000 description 2
- WNGVUZWBXZKQES-YUMQZZPRSA-N Leu-Ala-Gly Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](C)C(=O)NCC(O)=O WNGVUZWBXZKQES-YUMQZZPRSA-N 0.000 description 2
- 240000004658 Medicago sativa Species 0.000 description 2
- 235000017587 Medicago sativa ssp. sativa Nutrition 0.000 description 2
- OVBPIULPVIDEAO-UHFFFAOYSA-N N-Pteroyl-L-glutaminsaeure Natural products C=1N=C2NC(N)=NC(=O)C2=NC=1CNC1=CC=C(C(=O)NC(CCC(O)=O)C(O)=O)C=C1 OVBPIULPVIDEAO-UHFFFAOYSA-N 0.000 description 2
- 108010002311 N-glycylglutamic acid Proteins 0.000 description 2
- 108010065395 Neuropep-1 Proteins 0.000 description 2
- 108700026244 Open Reading Frames Proteins 0.000 description 2
- 240000007652 Petunia axillaris Species 0.000 description 2
- 235000004443 Ricinus communis Nutrition 0.000 description 2
- 241000736239 Ruellia Species 0.000 description 2
- NRCJWSGXMAPYQX-LPEHRKFASA-N Ser-Arg-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCCN=C(N)N)NC(=O)[C@H](CO)N)C(=O)O NRCJWSGXMAPYQX-LPEHRKFASA-N 0.000 description 2
- 239000005708 Sodium hypochlorite Substances 0.000 description 2
- OQWNEUXPKHIEJO-NRPADANISA-N Val-Glu-Ser Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CO)C(=O)O)N OQWNEUXPKHIEJO-NRPADANISA-N 0.000 description 2
- 241000700605 Viruses Species 0.000 description 2
- PTFCDOFLOPIGGS-UHFFFAOYSA-N Zinc dication Chemical compound [Zn+2] PTFCDOFLOPIGGS-UHFFFAOYSA-N 0.000 description 2
- 108010044940 alanylglutamine Proteins 0.000 description 2
- 230000003321 amplification Effects 0.000 description 2
- 230000015572 biosynthetic process Effects 0.000 description 2
- FPPNZSSZRUTDAP-UWFZAAFLSA-N carbenicillin Chemical compound N([C@H]1[C@H]2SC([C@@H](N2C1=O)C(O)=O)(C)C)C(=O)C(C(O)=O)C1=CC=CC=C1 FPPNZSSZRUTDAP-UWFZAAFLSA-N 0.000 description 2
- 229960003669 carbenicillin Drugs 0.000 description 2
- 239000012881 co-culture medium Substances 0.000 description 2
- 150000001875 compounds Chemical class 0.000 description 2
- 229910001431 copper ion Inorganic materials 0.000 description 2
- 238000010586 diagram Methods 0.000 description 2
- 238000004520 electroporation Methods 0.000 description 2
- 241001233957 eudicotyledons Species 0.000 description 2
- 210000003527 eukaryotic cell Anatomy 0.000 description 2
- 238000002474 experimental method Methods 0.000 description 2
- 229960000304 folic acid Drugs 0.000 description 2
- 235000019152 folic acid Nutrition 0.000 description 2
- 239000011724 folic acid Substances 0.000 description 2
- 108010063718 gamma-glutamylaspartic acid Proteins 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
- XKUKSGPZAADMRA-UHFFFAOYSA-N glycyl-glycyl-glycine Chemical compound NCC(=O)NCC(=O)NCC(O)=O XKUKSGPZAADMRA-UHFFFAOYSA-N 0.000 description 2
- 108010036413 histidylglycine Proteins 0.000 description 2
- 230000002779 inactivation Effects 0.000 description 2
- 238000011835 investigation Methods 0.000 description 2
- 108010064235 lysylglycine Proteins 0.000 description 2
- 229910021645 metal ion Inorganic materials 0.000 description 2
- 238000012986 modification Methods 0.000 description 2
- 230000004048 modification Effects 0.000 description 2
- 238000003199 nucleic acid amplification method Methods 0.000 description 2
- 230000036961 partial effect Effects 0.000 description 2
- 230000008569 process Effects 0.000 description 2
- 230000009467 reduction Effects 0.000 description 2
- 238000011160 research Methods 0.000 description 2
- 230000002441 reversible effect Effects 0.000 description 2
- 108010048818 seryl-histidine Proteins 0.000 description 2
- 239000011780 sodium chloride Substances 0.000 description 2
- SUKJFIGYRHOWBL-UHFFFAOYSA-N sodium hypochlorite Chemical compound [Na+].Cl[O-] SUKJFIGYRHOWBL-UHFFFAOYSA-N 0.000 description 2
- 238000003786 synthesis reaction Methods 0.000 description 2
- UZKQTCBAMSWPJD-UQCOIBPSSA-N trans-Zeatin Natural products OCC(/C)=C\CNC1=NC=NC2=C1N=CN2 UZKQTCBAMSWPJD-UQCOIBPSSA-N 0.000 description 2
- UZKQTCBAMSWPJD-FARCUNLSSA-N trans-zeatin Chemical compound OCC(/C)=C/CNC1=NC=NC2=C1N=CN2 UZKQTCBAMSWPJD-FARCUNLSSA-N 0.000 description 2
- 239000012137 tryptone Substances 0.000 description 2
- 229940023877 zeatin Drugs 0.000 description 2
- JNTMAZFVYNDPLB-PEDHHIEDSA-N (2S,3S)-2-[[[(2S)-1-[(2S,3S)-2-amino-3-methyl-1-oxopentyl]-2-pyrrolidinyl]-oxomethyl]amino]-3-methylpentanoic acid Chemical compound CC[C@H](C)[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)CC)C(O)=O JNTMAZFVYNDPLB-PEDHHIEDSA-N 0.000 description 1
- 239000005971 1-naphthylacetic acid Substances 0.000 description 1
- KDZIGQIDPXKMBA-UHFFFAOYSA-N 2-[[2-[[2-[(2-amino-3-methylbutanoyl)amino]acetyl]amino]-3-hydroxypropanoyl]amino]pentanedioic acid Chemical compound CC(C)C(N)C(=O)NCC(=O)NC(CO)C(=O)NC(C(O)=O)CCC(O)=O KDZIGQIDPXKMBA-UHFFFAOYSA-N 0.000 description 1
- IIDAJRNSZSFFCB-UHFFFAOYSA-N 4-amino-5-methoxy-2-methylbenzenesulfonamide Chemical compound COC1=CC(S(N)(=O)=O)=C(C)C=C1N IIDAJRNSZSFFCB-UHFFFAOYSA-N 0.000 description 1
- 239000005972 6-Benzyladenine Substances 0.000 description 1
- SBGXWWCLHIOABR-UHFFFAOYSA-N Ala Ala Gly Ala Chemical compound CC(N)C(=O)NC(C)C(=O)NCC(=O)NC(C)C(O)=O SBGXWWCLHIOABR-UHFFFAOYSA-N 0.000 description 1
- DKJPOZOEBONHFS-ZLUOBGJFSA-N Ala-Ala-Asp Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CC(O)=O DKJPOZOEBONHFS-ZLUOBGJFSA-N 0.000 description 1
- YLTKNGYYPIWKHZ-ACZMJKKPSA-N Ala-Ala-Glu Chemical compound C[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCC(O)=O YLTKNGYYPIWKHZ-ACZMJKKPSA-N 0.000 description 1
- JBVSSSZFNTXJDX-YTLHQDLWSA-N Ala-Ala-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@H](C)N JBVSSSZFNTXJDX-YTLHQDLWSA-N 0.000 description 1
- SVBXIUDNTRTKHE-CIUDSAMLSA-N Ala-Arg-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(O)=O)C(O)=O SVBXIUDNTRTKHE-CIUDSAMLSA-N 0.000 description 1
- YAXNATKKPOWVCP-ZLUOBGJFSA-N Ala-Asn-Ala Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](C)C(O)=O YAXNATKKPOWVCP-ZLUOBGJFSA-N 0.000 description 1
- NHCPCLJZRSIDHS-ZLUOBGJFSA-N Ala-Asp-Ala Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C)C(O)=O NHCPCLJZRSIDHS-ZLUOBGJFSA-N 0.000 description 1
- DECCMEWNXSNSDO-ZLUOBGJFSA-N Ala-Cys-Ala Chemical compound C[C@H](N)C(=O)N[C@@H](CS)C(=O)N[C@@H](C)C(O)=O DECCMEWNXSNSDO-ZLUOBGJFSA-N 0.000 description 1
- CRWFEKLFPVRPBV-CIUDSAMLSA-N Ala-Gln-Met Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCSC)C(O)=O CRWFEKLFPVRPBV-CIUDSAMLSA-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
- HMRWQTHUDVXMGH-GUBZILKMSA-N Ala-Glu-Lys Chemical compound C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CCCCN HMRWQTHUDVXMGH-GUBZILKMSA-N 0.000 description 1
- HQJKCXHQNUCKMY-GHCJXIJMSA-N Ala-Ile-Asp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)[C@H](C)N HQJKCXHQNUCKMY-GHCJXIJMSA-N 0.000 description 1
- CKLDHDOIYBVUNP-KBIXCLLPSA-N Ala-Ile-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCC(O)=O)C(O)=O CKLDHDOIYBVUNP-KBIXCLLPSA-N 0.000 description 1
- VNYMOTCMNHJGTG-JBDRJPRFSA-N Ala-Ile-Ser Chemical compound [H]N[C@@H](C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CO)C(O)=O VNYMOTCMNHJGTG-JBDRJPRFSA-N 0.000 description 1
- LNNSWWRRYJLGNI-NAKRPEOUSA-N Ala-Ile-Val Chemical compound C[C@H](N)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C(C)C)C(O)=O LNNSWWRRYJLGNI-NAKRPEOUSA-N 0.000 description 1
- OYJCVIGKMXUVKB-GARJFASQSA-N Ala-Leu-Pro Chemical compound C[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)N1CCC[C@@H]1C(=O)O)N OYJCVIGKMXUVKB-GARJFASQSA-N 0.000 description 1
- QUIGLPSHIFPEOV-CIUDSAMLSA-N Ala-Lys-Ala Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O QUIGLPSHIFPEOV-CIUDSAMLSA-N 0.000 description 1
- SUHLZMHFRALVSY-YUMQZZPRSA-N Ala-Lys-Gly Chemical compound NCCCC[C@H](NC(=O)[C@@H](N)C)C(=O)NCC(O)=O SUHLZMHFRALVSY-YUMQZZPRSA-N 0.000 description 1
- VCSABYLVNWQYQE-UHFFFAOYSA-N Ala-Lys-Lys Natural products NCCCCC(NC(=O)C(N)C)C(=O)NC(CCCCN)C(O)=O VCSABYLVNWQYQE-UHFFFAOYSA-N 0.000 description 1
- CHFFHQUVXHEGBY-GARJFASQSA-N Ala-Lys-Pro Chemical compound C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N1CCC[C@@H]1C(=O)O)N CHFFHQUVXHEGBY-GARJFASQSA-N 0.000 description 1
- XSTZMVAYYCJTNR-DCAQKATOSA-N Ala-Met-Leu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O XSTZMVAYYCJTNR-DCAQKATOSA-N 0.000 description 1
- VJVQKGYHIZPSNS-FXQIFTODSA-N Ala-Ser-Arg Chemical compound C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCCN=C(N)N VJVQKGYHIZPSNS-FXQIFTODSA-N 0.000 description 1
- IOFVWPYSRSCWHI-JXUBOQSCSA-N Ala-Thr-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H]([C@@H](C)O)NC(=O)[C@H](C)N IOFVWPYSRSCWHI-JXUBOQSCSA-N 0.000 description 1
- QOIGKCBMXUCDQU-KDXUFGMBSA-N Ala-Thr-Pro Chemical compound C[C@H]([C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](C)N)O QOIGKCBMXUCDQU-KDXUFGMBSA-N 0.000 description 1
- VQBULXOHAZSTQY-GKCIPKSASA-N Ala-Trp-Phe Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O VQBULXOHAZSTQY-GKCIPKSASA-N 0.000 description 1
- ZXKNLCPUNZPFGY-LEWSCRJBSA-N Ala-Tyr-Pro Chemical compound C[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N2CCC[C@@H]2C(=O)O)N ZXKNLCPUNZPFGY-LEWSCRJBSA-N 0.000 description 1
- XKHLBBQNPSOGPI-GUBZILKMSA-N Ala-Val-Met Chemical compound CSCC[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@H](C)N XKHLBBQNPSOGPI-GUBZILKMSA-N 0.000 description 1
- OTOXOKCIIQLMFH-KZVJFYERSA-N Arg-Ala-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@@H](N)CCCN=C(N)N OTOXOKCIIQLMFH-KZVJFYERSA-N 0.000 description 1
- TTXYKSADPSNOIF-IHRRRGAJSA-N Arg-Asp-Phe Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O TTXYKSADPSNOIF-IHRRRGAJSA-N 0.000 description 1
- XLWSGICNBZGYTA-CIUDSAMLSA-N Arg-Glu-Asp Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O XLWSGICNBZGYTA-CIUDSAMLSA-N 0.000 description 1
- SKTGPBFTMNLIHQ-KKUMJFAQSA-N Arg-Glu-Phe Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O SKTGPBFTMNLIHQ-KKUMJFAQSA-N 0.000 description 1
- NVUIWHJLPSZZQC-CYDGBPFRSA-N Arg-Ile-Arg Chemical compound NC(N)=NCCC[C@H](N)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O NVUIWHJLPSZZQC-CYDGBPFRSA-N 0.000 description 1
- FFEUXEAKYRCACT-PEDHHIEDSA-N Arg-Ile-Ile Chemical compound CC[C@H](C)[C@H](NC(=O)[C@@H](NC(=O)[C@@H](N)CCCNC(N)=N)[C@@H](C)CC)C(O)=O FFEUXEAKYRCACT-PEDHHIEDSA-N 0.000 description 1
- OOIMKQRCPJBGPD-XUXIUFHCSA-N Arg-Ile-Leu Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(C)C)C(O)=O OOIMKQRCPJBGPD-XUXIUFHCSA-N 0.000 description 1
- LKDHUGLXOHYINY-XUXIUFHCSA-N Arg-Ile-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N LKDHUGLXOHYINY-XUXIUFHCSA-N 0.000 description 1
- LLUGJARLJCGLAR-CYDGBPFRSA-N Arg-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](CCCN=C(N)N)N LLUGJARLJCGLAR-CYDGBPFRSA-N 0.000 description 1
- NGTYEHIRESTSRX-UWVGGRQHSA-N Arg-Lys-Gly Chemical compound NCCCC[C@@H](C(=O)NCC(O)=O)NC(=O)[C@@H](N)CCCN=C(N)N NGTYEHIRESTSRX-UWVGGRQHSA-N 0.000 description 1
- UIUXXFIKWQVMEX-UFYCRDLUSA-N Arg-Phe-Tyr Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O UIUXXFIKWQVMEX-UFYCRDLUSA-N 0.000 description 1
- JPAWCMXVNZPJLO-IHRRRGAJSA-N Arg-Ser-Phe Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O JPAWCMXVNZPJLO-IHRRRGAJSA-N 0.000 description 1
- YNDLOUMBVDVALC-ZLUOBGJFSA-N Asn-Ala-Ala Chemical compound C[C@@H](C(=O)N[C@@H](C)C(=O)O)NC(=O)[C@H](CC(=O)N)N YNDLOUMBVDVALC-ZLUOBGJFSA-N 0.000 description 1
- VDCIPFYVCICPEC-FXQIFTODSA-N Asn-Arg-Ala Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](C)C(O)=O VDCIPFYVCICPEC-FXQIFTODSA-N 0.000 description 1
- ZMWDUIIACVLIHK-GHCJXIJMSA-N Asn-Cys-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CS)NC(=O)[C@H](CC(=O)N)N ZMWDUIIACVLIHK-GHCJXIJMSA-N 0.000 description 1
- BZMWJLLUAKSIMH-FXQIFTODSA-N Asn-Glu-Glu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O BZMWJLLUAKSIMH-FXQIFTODSA-N 0.000 description 1
- OOWSBIOUKIUWLO-RCOVLWMOSA-N Asn-Gly-Val Chemical compound [H]N[C@@H](CC(N)=O)C(=O)NCC(=O)N[C@@H](C(C)C)C(O)=O OOWSBIOUKIUWLO-RCOVLWMOSA-N 0.000 description 1
- FHETWELNCBMRMG-HJGDQZAQSA-N Asn-Leu-Thr Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O FHETWELNCBMRMG-HJGDQZAQSA-N 0.000 description 1
- COWITDLVHMZSIW-CIUDSAMLSA-N Asn-Lys-Ser Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(O)=O COWITDLVHMZSIW-CIUDSAMLSA-N 0.000 description 1
- AEZCCDMZZJOGII-DCAQKATOSA-N Asn-Met-Leu Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O AEZCCDMZZJOGII-DCAQKATOSA-N 0.000 description 1
- QIRJQYQOIKBPBZ-IHRRRGAJSA-N Asn-Tyr-Arg Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O QIRJQYQOIKBPBZ-IHRRRGAJSA-N 0.000 description 1
- WQAOZCVOOYUWKG-LSJOCFKGSA-N Asn-Val-Val Chemical compound CC(C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)O)NC(=O)[C@H](CC(=O)N)N WQAOZCVOOYUWKG-LSJOCFKGSA-N 0.000 description 1
- UGKZHCBLMLSANF-CIUDSAMLSA-N Asp-Asn-Leu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(C)C)C(O)=O UGKZHCBLMLSANF-CIUDSAMLSA-N 0.000 description 1
- GHODABZPVZMWCE-FXQIFTODSA-N Asp-Glu-Glu Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O GHODABZPVZMWCE-FXQIFTODSA-N 0.000 description 1
- VFUXXFVCYZPOQG-WDSKDSINSA-N Asp-Glu-Gly Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)NCC(O)=O VFUXXFVCYZPOQG-WDSKDSINSA-N 0.000 description 1
- YDJVIBMKAMQPPP-LAEOZQHASA-N Asp-Glu-Val Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O YDJVIBMKAMQPPP-LAEOZQHASA-N 0.000 description 1
- PSLSTUMPZILTAH-BYULHYEWSA-N Asp-Gly-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CC(O)=O PSLSTUMPZILTAH-BYULHYEWSA-N 0.000 description 1
- KHGPWGKPYHPOIK-QWRGUYRKSA-N Asp-Gly-Phe Chemical compound [H]N[C@@H](CC(O)=O)C(=O)NCC(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O KHGPWGKPYHPOIK-QWRGUYRKSA-N 0.000 description 1
- WSGVTKZFVJSJOG-RCOVLWMOSA-N Asp-Gly-Val Chemical compound [H]N[C@@H](CC(O)=O)C(=O)NCC(=O)N[C@@H](C(C)C)C(O)=O WSGVTKZFVJSJOG-RCOVLWMOSA-N 0.000 description 1
- TZOZNVLBTAFJRW-UGYAYLCHSA-N Asp-Ile-Asp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)O)NC(=O)[C@H](CC(=O)O)N TZOZNVLBTAFJRW-UGYAYLCHSA-N 0.000 description 1
- KYQNAIMCTRZLNP-QSFUFRPTSA-N Asp-Ile-Val Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C(C)C)C(O)=O KYQNAIMCTRZLNP-QSFUFRPTSA-N 0.000 description 1
- GWIJZUVQVDJHDI-AVGNSLFASA-N Asp-Phe-Glu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(O)=O)C(O)=O GWIJZUVQVDJHDI-AVGNSLFASA-N 0.000 description 1
- UCHSVZYJKJLPHF-BZSNNMDCSA-N Asp-Phe-Phe Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O UCHSVZYJKJLPHF-BZSNNMDCSA-N 0.000 description 1
- RPUYTJJZXQBWDT-SRVKXCTJSA-N Asp-Phe-Ser Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](CC(=O)O)N RPUYTJJZXQBWDT-SRVKXCTJSA-N 0.000 description 1
- DINOVZWPTMGSRF-QXEWZRGKSA-N Asp-Pro-Val Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(O)=O DINOVZWPTMGSRF-QXEWZRGKSA-N 0.000 description 1
- OFYVKOXTTDCUIL-FXQIFTODSA-N Asp-Ser-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CC(=O)O)N OFYVKOXTTDCUIL-FXQIFTODSA-N 0.000 description 1
- UXRVDHVARNBOIO-QSFUFRPTSA-N Asp-Val-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](C(C)C)NC(=O)[C@H](CC(=O)O)N UXRVDHVARNBOIO-QSFUFRPTSA-N 0.000 description 1
- 239000002028 Biomass Substances 0.000 description 1
- 102000020856 Copper Transport Proteins Human genes 0.000 description 1
- 108091004554 Copper Transport Proteins Proteins 0.000 description 1
- LHLSSZYQFUNWRZ-NAKRPEOUSA-N Cys-Arg-Ile Chemical compound [H]N[C@@H](CS)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O LHLSSZYQFUNWRZ-NAKRPEOUSA-N 0.000 description 1
- WTXCNOPZMQRTNN-BWBBJGPYSA-N Cys-Thr-Ser Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](CS)N)O WTXCNOPZMQRTNN-BWBBJGPYSA-N 0.000 description 1
- 230000006820 DNA synthesis Effects 0.000 description 1
- 241000588724 Escherichia coli Species 0.000 description 1
- VTLYFUHAOXGGBS-UHFFFAOYSA-N Fe3+ Chemical compound [Fe+3] VTLYFUHAOXGGBS-UHFFFAOYSA-N 0.000 description 1
- 108091092584 GDNA Proteins 0.000 description 1
- KVYVOGYEMPEXBT-GUBZILKMSA-N Gln-Ala-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](C)NC(=O)[C@@H](N)CCC(N)=O KVYVOGYEMPEXBT-GUBZILKMSA-N 0.000 description 1
- XXLBHPPXDUWYAG-XQXXSGGOSA-N Gln-Ala-Thr Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O XXLBHPPXDUWYAG-XQXXSGGOSA-N 0.000 description 1
- GNMQDOGFWYWPNM-LAEOZQHASA-N Gln-Gly-Ile Chemical compound CC[C@H](C)[C@H](NC(=O)CNC(=O)[C@@H](N)CCC(N)=O)C(O)=O GNMQDOGFWYWPNM-LAEOZQHASA-N 0.000 description 1
- XFAUJGNLHIGXET-AVGNSLFASA-N Gln-Leu-Leu Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O XFAUJGNLHIGXET-AVGNSLFASA-N 0.000 description 1
- JNENSVNAUWONEZ-GUBZILKMSA-N Gln-Lys-Asn Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(N)=O)C(O)=O JNENSVNAUWONEZ-GUBZILKMSA-N 0.000 description 1
- QKWBEMCLYTYBNI-GVXVVHGQSA-N Gln-Lys-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CCC(N)=O QKWBEMCLYTYBNI-GVXVVHGQSA-N 0.000 description 1
- WHVLABLIJYGVEK-QEWYBTABSA-N Gln-Phe-Ile Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O WHVLABLIJYGVEK-QEWYBTABSA-N 0.000 description 1
- JILRMFFFCHUUTJ-ACZMJKKPSA-N Gln-Ser-Ser Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CO)C(O)=O JILRMFFFCHUUTJ-ACZMJKKPSA-N 0.000 description 1
- SDSMVVSHLAAOJL-UKJIMTQDSA-N Gln-Val-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](C(C)C)NC(=O)[C@H](CCC(=O)N)N SDSMVVSHLAAOJL-UKJIMTQDSA-N 0.000 description 1
- RLZBLVSJDFHDBL-KBIXCLLPSA-N Glu-Ala-Ile Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O RLZBLVSJDFHDBL-KBIXCLLPSA-N 0.000 description 1
- VAZZOGXDUQSVQF-NUMRIWBASA-N Glu-Asn-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CC(=O)N)NC(=O)[C@H](CCC(=O)O)N)O VAZZOGXDUQSVQF-NUMRIWBASA-N 0.000 description 1
- QPRZKNOOOBWXSU-CIUDSAMLSA-N Glu-Asp-Arg Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CCCN=C(N)N QPRZKNOOOBWXSU-CIUDSAMLSA-N 0.000 description 1
- HJIFPJUEOGZWRI-GUBZILKMSA-N Glu-Asp-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H](CCC(=O)O)N HJIFPJUEOGZWRI-GUBZILKMSA-N 0.000 description 1
- WPLGNDORMXTMQS-FXQIFTODSA-N Glu-Gln-Ser Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(O)=O WPLGNDORMXTMQS-FXQIFTODSA-N 0.000 description 1
- QQLBPVKLJBAXBS-FXQIFTODSA-N Glu-Glu-Asn Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O QQLBPVKLJBAXBS-FXQIFTODSA-N 0.000 description 1
- KASDBWKLWJKTLJ-GUBZILKMSA-N Glu-Glu-Met Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCSC)C(O)=O KASDBWKLWJKTLJ-GUBZILKMSA-N 0.000 description 1
- VGOFRWOTSXVPAU-SDDRHHMPSA-N Glu-His-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC2=CN=CN2)NC(=O)[C@H](CCC(=O)O)N)C(=O)O VGOFRWOTSXVPAU-SDDRHHMPSA-N 0.000 description 1
- LGYCLOCORAEQSZ-PEFMBERDSA-N Glu-Ile-Asp Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC(O)=O)C(O)=O LGYCLOCORAEQSZ-PEFMBERDSA-N 0.000 description 1
- ITBHUUMCJJQUSC-LAEOZQHASA-N Glu-Ile-Gly Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)NCC(O)=O ITBHUUMCJJQUSC-LAEOZQHASA-N 0.000 description 1
- IVGJYOOGJLFKQE-AVGNSLFASA-N Glu-Leu-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CCC(=O)O)N IVGJYOOGJLFKQE-AVGNSLFASA-N 0.000 description 1
- OQXDUSZKISQQSS-GUBZILKMSA-N Glu-Lys-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O OQXDUSZKISQQSS-GUBZILKMSA-N 0.000 description 1
- BPLNJYHNAJVLRT-ACZMJKKPSA-N Glu-Ser-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](C)C(O)=O BPLNJYHNAJVLRT-ACZMJKKPSA-N 0.000 description 1
- GUOWMVFLAJNPDY-CIUDSAMLSA-N Glu-Ser-Met Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCSC)C(O)=O GUOWMVFLAJNPDY-CIUDSAMLSA-N 0.000 description 1
- RGJKYNUINKGPJN-RWRJDSDZSA-N Glu-Thr-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H]([C@@H](C)O)NC(=O)[C@H](CCC(=O)O)N RGJKYNUINKGPJN-RWRJDSDZSA-N 0.000 description 1
- MXJYXYDREQWUMS-XKBZYTNZSA-N Glu-Thr-Ser Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(O)=O MXJYXYDREQWUMS-XKBZYTNZSA-N 0.000 description 1
- HVKAAUOFFTUSAA-XDTLVQLUSA-N Glu-Tyr-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](C)C(O)=O HVKAAUOFFTUSAA-XDTLVQLUSA-N 0.000 description 1
- HQTDNEZTGZUWSY-XVKPBYJWSA-N Glu-Val-Gly Chemical compound CC(C)[C@H](NC(=O)[C@@H](N)CCC(O)=O)C(=O)NCC(O)=O HQTDNEZTGZUWSY-XVKPBYJWSA-N 0.000 description 1
- VIPDPMHGICREIS-GVXVVHGQSA-N Glu-Val-Leu Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O VIPDPMHGICREIS-GVXVVHGQSA-N 0.000 description 1
- UGVQELHRNUDMAA-BYPYZUCNSA-N Gly-Ala-Gly Chemical compound [NH3+]CC(=O)N[C@@H](C)C(=O)NCC([O-])=O UGVQELHRNUDMAA-BYPYZUCNSA-N 0.000 description 1
- XQHSBNVACKQWAV-WHFBIAKZSA-N Gly-Asp-Asn Chemical compound [H]NCC(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(N)=O)C(O)=O XQHSBNVACKQWAV-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
- GNPVTZJUUBPZKW-WDSKDSINSA-N Gly-Gln-Ser Chemical compound [H]NCC(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(O)=O GNPVTZJUUBPZKW-WDSKDSINSA-N 0.000 description 1
- UQJNXZSSGQIPIQ-FBCQKBJTSA-N Gly-Gly-Thr Chemical compound C[C@@H](O)[C@@H](C(O)=O)NC(=O)CNC(=O)CN UQJNXZSSGQIPIQ-FBCQKBJTSA-N 0.000 description 1
- ADZGCWWDPFDHCY-ZETCQYMHSA-N Gly-His-Gly Chemical compound OC(=O)CNC(=O)[C@@H](NC(=O)CN)CC1=CN=CN1 ADZGCWWDPFDHCY-ZETCQYMHSA-N 0.000 description 1
- HMHRTKOWRUPPNU-RCOVLWMOSA-N Gly-Ile-Gly Chemical compound NCC(=O)N[C@@H]([C@@H](C)CC)C(=O)NCC(O)=O HMHRTKOWRUPPNU-RCOVLWMOSA-N 0.000 description 1
- SCWYHUQOOFRVHP-MBLNEYKQSA-N Gly-Ile-Thr Chemical compound NCC(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O SCWYHUQOOFRVHP-MBLNEYKQSA-N 0.000 description 1
- WDEHMRNSGHVNOH-VHSXEESVSA-N Gly-Lys-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CCCCN)NC(=O)CN)C(=O)O WDEHMRNSGHVNOH-VHSXEESVSA-N 0.000 description 1
- OQQKUTVULYLCDG-ONGXEEELSA-N Gly-Lys-Val Chemical compound CC(C)[C@H](NC(=O)[C@H](CCCCN)NC(=O)CN)C(O)=O OQQKUTVULYLCDG-ONGXEEELSA-N 0.000 description 1
- UWQDKRIZSROAKS-FJXKBIBVSA-N Gly-Met-Thr Chemical compound [H]NCC(=O)N[C@@H](CCSC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O UWQDKRIZSROAKS-FJXKBIBVSA-N 0.000 description 1
- IRJWAYCXIYUHQE-WHFBIAKZSA-N Gly-Ser-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CO)NC(=O)CN IRJWAYCXIYUHQE-WHFBIAKZSA-N 0.000 description 1
- ZLCLYFGMKFCDCN-XPUUQOCRSA-N Gly-Ser-Val Chemical compound CC(C)[C@H](NC(=O)[C@H](CO)NC(=O)CN)C(O)=O ZLCLYFGMKFCDCN-XPUUQOCRSA-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
- NGRPGJGKJMUGDM-XVKPBYJWSA-N Gly-Val-Gln Chemical compound NCC(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O NGRPGJGKJMUGDM-XVKPBYJWSA-N 0.000 description 1
- SYOJVRNQCXYEOV-XVKPBYJWSA-N Gly-Val-Glu Chemical compound [H]NCC(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O SYOJVRNQCXYEOV-XVKPBYJWSA-N 0.000 description 1
- RYAOJUMWLWUGNW-QMMMGPOBSA-N Gly-Val-Gly Chemical compound NCC(=O)N[C@@H](C(C)C)C(=O)NCC(O)=O RYAOJUMWLWUGNW-QMMMGPOBSA-N 0.000 description 1
- BAYQNCWLXIDLHX-ONGXEEELSA-N Gly-Val-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)CN BAYQNCWLXIDLHX-ONGXEEELSA-N 0.000 description 1
- AFMOTCMSEBITOE-YEPSODPASA-N Gly-Val-Thr Chemical compound NCC(=O)N[C@@H](C(C)C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O AFMOTCMSEBITOE-YEPSODPASA-N 0.000 description 1
- HDXNWVLQSQFJOX-SRVKXCTJSA-N His-Arg-Gln Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N HDXNWVLQSQFJOX-SRVKXCTJSA-N 0.000 description 1
- MDOBWSFNSNPENN-PMVVWTBXSA-N His-Thr-Gly Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H]([C@@H](C)O)C(=O)NCC(O)=O MDOBWSFNSNPENN-PMVVWTBXSA-N 0.000 description 1
- ISQOVWDWRUONJH-YESZJQIVSA-N His-Tyr-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC2=CC=C(C=C2)O)NC(=O)[C@H](CC3=CN=CN3)N)C(=O)O ISQOVWDWRUONJH-YESZJQIVSA-N 0.000 description 1
- GYXDQXPCPASCNR-NHCYSSNCSA-N His-Val-Asn Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CC1=CN=CN1)N GYXDQXPCPASCNR-NHCYSSNCSA-N 0.000 description 1
- CISBRYJZMFWOHJ-JBDRJPRFSA-N Ile-Ala-Cys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](C)C(=O)N[C@@H](CS)C(=O)O)N CISBRYJZMFWOHJ-JBDRJPRFSA-N 0.000 description 1
- 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 1
- RWIKBYVJQAJYDP-BJDJZHNGSA-N Ile-Ala-Lys Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@H](C(O)=O)CCCCN RWIKBYVJQAJYDP-BJDJZHNGSA-N 0.000 description 1
- TZCGZYWNIDZZMR-UHFFFAOYSA-N Ile-Arg-Ala Natural products CCC(C)C(N)C(=O)NC(C(=O)NC(C)C(O)=O)CCCN=C(N)N TZCGZYWNIDZZMR-UHFFFAOYSA-N 0.000 description 1
- FVEWRQXNISSYFO-ZPFDUUQYSA-N Ile-Arg-Glu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N FVEWRQXNISSYFO-ZPFDUUQYSA-N 0.000 description 1
- AZEYWPUCOYXFOE-CYDGBPFRSA-N Ile-Arg-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](C(C)C)C(=O)O)N AZEYWPUCOYXFOE-CYDGBPFRSA-N 0.000 description 1
- NPROWIBAWYMPAZ-GUDRVLHUSA-N Ile-Asp-Pro Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N NPROWIBAWYMPAZ-GUDRVLHUSA-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
- NHJKZMDIMMTVCK-QXEWZRGKSA-N Ile-Gly-Arg Chemical compound CC[C@H](C)[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCN=C(N)N NHJKZMDIMMTVCK-QXEWZRGKSA-N 0.000 description 1
- YNMQUIVKEFRCPH-QSFUFRPTSA-N Ile-Ile-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)CC)C(=O)NCC(=O)O)N YNMQUIVKEFRCPH-QSFUFRPTSA-N 0.000 description 1
- GAZGFPOZOLEYAJ-YTFOTSKYSA-N Ile-Leu-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)O)N GAZGFPOZOLEYAJ-YTFOTSKYSA-N 0.000 description 1
- HPCFRQWLTRDGHT-AJNGGQMLSA-N Ile-Leu-Leu Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O HPCFRQWLTRDGHT-AJNGGQMLSA-N 0.000 description 1
- NZGTYCMLUGYMCV-XUXIUFHCSA-N Ile-Lys-Arg Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)N NZGTYCMLUGYMCV-XUXIUFHCSA-N 0.000 description 1
- KCTIFOCXAIUQQK-QXEWZRGKSA-N Ile-Pro-Gly Chemical compound CC[C@H](C)[C@H](N)C(=O)N1CCC[C@H]1C(=O)NCC(O)=O KCTIFOCXAIUQQK-QXEWZRGKSA-N 0.000 description 1
- MLSUZXHSNRBDCI-CYDGBPFRSA-N Ile-Pro-Val Chemical compound CC[C@H](C)[C@@H](C(=O)N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(=O)O)N MLSUZXHSNRBDCI-CYDGBPFRSA-N 0.000 description 1
- ZDNNDIJTUHQCAM-MXAVVETBSA-N Ile-Ser-Phe Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)N ZDNNDIJTUHQCAM-MXAVVETBSA-N 0.000 description 1
- RQJUKVXWAKJDBW-SVSWQMSJSA-N Ile-Ser-Thr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(=O)O)N RQJUKVXWAKJDBW-SVSWQMSJSA-N 0.000 description 1
- ZGKVPOSSTGHJAF-HJPIBITLSA-N Ile-Tyr-Ser Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)N[C@@H](CO)C(=O)O)N ZGKVPOSSTGHJAF-HJPIBITLSA-N 0.000 description 1
- KXUKTDGKLAOCQK-LSJOCFKGSA-N Ile-Val-Gly Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)NCC(O)=O KXUKTDGKLAOCQK-LSJOCFKGSA-N 0.000 description 1
- UYODHPPSCXBNCS-XUXIUFHCSA-N Ile-Val-Leu Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CC(C)C UYODHPPSCXBNCS-XUXIUFHCSA-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
- 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
- KWTVLKBOQATPHJ-SRVKXCTJSA-N Leu-Ala-Lys Chemical compound C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CC(C)C)N KWTVLKBOQATPHJ-SRVKXCTJSA-N 0.000 description 1
- XBBKIIGCUMBKCO-JXUBOQSCSA-N Leu-Ala-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](C)C(=O)N[C@@H]([C@@H](C)O)C(O)=O XBBKIIGCUMBKCO-JXUBOQSCSA-N 0.000 description 1
- FJUKMPUELVROGK-IHRRRGAJSA-N Leu-Arg-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N FJUKMPUELVROGK-IHRRRGAJSA-N 0.000 description 1
- XYUBOFCTGPZFSA-WDSOQIARSA-N Leu-Arg-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@@H](N)CC(C)C)C(O)=O)=CNC2=C1 XYUBOFCTGPZFSA-WDSOQIARSA-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
- KTFHTMHHKXUYPW-ZPFDUUQYSA-N Leu-Asp-Ile Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O KTFHTMHHKXUYPW-ZPFDUUQYSA-N 0.000 description 1
- BOFAFKVZQUMTID-AVGNSLFASA-N Leu-Gln-His Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)N BOFAFKVZQUMTID-AVGNSLFASA-N 0.000 description 1
- HFBCHNRFRYLZNV-GUBZILKMSA-N Leu-Glu-Asp Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O HFBCHNRFRYLZNV-GUBZILKMSA-N 0.000 description 1
- HYIFFZAQXPUEAU-QWRGUYRKSA-N Leu-Gly-Leu Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CC(C)C HYIFFZAQXPUEAU-QWRGUYRKSA-N 0.000 description 1
- HYMLKESRWLZDBR-WEDXCCLWSA-N Leu-Gly-Thr Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(O)=O HYMLKESRWLZDBR-WEDXCCLWSA-N 0.000 description 1
- XQXGNBFMAXWIGI-MXAVVETBSA-N Leu-His-Ile Chemical compound CC[C@H](C)[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)CC(C)C)CC1=CN=CN1 XQXGNBFMAXWIGI-MXAVVETBSA-N 0.000 description 1
- DBSLVQBXKVKDKJ-BJDJZHNGSA-N Leu-Ile-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C)C(O)=O DBSLVQBXKVKDKJ-BJDJZHNGSA-N 0.000 description 1
- OMHLATXVNQSALM-FQUUOJAGSA-N Leu-Ile-Pro Chemical compound CC[C@H](C)[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CC(C)C)N OMHLATXVNQSALM-FQUUOJAGSA-N 0.000 description 1
- LIINDKYIGYTDLG-PPCPHDFISA-N Leu-Ile-Thr Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O LIINDKYIGYTDLG-PPCPHDFISA-N 0.000 description 1
- YOKVEHGYYQEQOP-QWRGUYRKSA-N Leu-Leu-Gly Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)NCC(O)=O YOKVEHGYYQEQOP-QWRGUYRKSA-N 0.000 description 1
- LXKNSJLSGPNHSK-KKUMJFAQSA-N Leu-Leu-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)O)N LXKNSJLSGPNHSK-KKUMJFAQSA-N 0.000 description 1
- WXUOJXIGOPMDJM-SRVKXCTJSA-N Leu-Lys-Asn Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(N)=O)C(O)=O WXUOJXIGOPMDJM-SRVKXCTJSA-N 0.000 description 1
- FKQPWMZLIIATBA-AJNGGQMLSA-N Leu-Lys-Ile Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O FKQPWMZLIIATBA-AJNGGQMLSA-N 0.000 description 1
- DDVHDMSBLRAKNV-IHRRRGAJSA-N Leu-Met-Leu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O DDVHDMSBLRAKNV-IHRRRGAJSA-N 0.000 description 1
- MJWVXZABPOKJJF-ACRUOGEOSA-N Leu-Phe-Phe 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=CC=C1)C(O)=O MJWVXZABPOKJJF-ACRUOGEOSA-N 0.000 description 1
- UHNQRAFSEBGZFZ-YESZJQIVSA-N Leu-Phe-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N2CCC[C@@H]2C(=O)O)N UHNQRAFSEBGZFZ-YESZJQIVSA-N 0.000 description 1
- IDGZVZJLYFTXSL-DCAQKATOSA-N Leu-Ser-Arg Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCCN=C(N)N IDGZVZJLYFTXSL-DCAQKATOSA-N 0.000 description 1
- URHJPNHRQMQGOZ-RHYQMDGZSA-N Leu-Thr-Met Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCSC)C(O)=O URHJPNHRQMQGOZ-RHYQMDGZSA-N 0.000 description 1
- WBRJVRXEGQIDRK-XIRDDKMYSA-N Leu-Trp-Ser Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@@H](N)CC(C)C)C(=O)N[C@@H](CO)C(O)=O)=CNC2=C1 WBRJVRXEGQIDRK-XIRDDKMYSA-N 0.000 description 1
- AIMGJYMCTAABEN-GVXVVHGQSA-N Leu-Val-Glu Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O AIMGJYMCTAABEN-GVXVVHGQSA-N 0.000 description 1
- 241000209510 Liliopsida Species 0.000 description 1
- 108010062166 Lys-Asn-Asp Proteins 0.000 description 1
- AAORVPFVUIHEAB-YUMQZZPRSA-N Lys-Asp-Gly Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(O)=O)C(=O)NCC(O)=O AAORVPFVUIHEAB-YUMQZZPRSA-N 0.000 description 1
- GJJQCBVRWDGLMQ-GUBZILKMSA-N Lys-Glu-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O GJJQCBVRWDGLMQ-GUBZILKMSA-N 0.000 description 1
- 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
- ISHNZELVUVPCHY-ZETCQYMHSA-N Lys-Gly-Gly Chemical compound NCCCC[C@H](N)C(=O)NCC(=O)NCC(O)=O ISHNZELVUVPCHY-ZETCQYMHSA-N 0.000 description 1
- ORVFEGYUJITPGI-IHRRRGAJSA-N Lys-Leu-Met Chemical compound CSCC[C@@H](C(O)=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H](N)CCCCN ORVFEGYUJITPGI-IHRRRGAJSA-N 0.000 description 1
- WRODMZBHNNPRLN-SRVKXCTJSA-N Lys-Leu-Ser Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O WRODMZBHNNPRLN-SRVKXCTJSA-N 0.000 description 1
- LJADEBULDNKJNK-IHRRRGAJSA-N Lys-Leu-Val Chemical compound CC(C)C[C@H](NC(=O)[C@@H](N)CCCCN)C(=O)N[C@@H](C(C)C)C(O)=O LJADEBULDNKJNK-IHRRRGAJSA-N 0.000 description 1
- HVAUKHLDSDDROB-KKUMJFAQSA-N Lys-Lys-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(O)=O HVAUKHLDSDDROB-KKUMJFAQSA-N 0.000 description 1
- ATNKHRAIZCMCCN-BZSNNMDCSA-N Lys-Lys-Phe Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](CCCCN)N ATNKHRAIZCMCCN-BZSNNMDCSA-N 0.000 description 1
- QQPSCXKFDSORFT-IHRRRGAJSA-N Lys-Lys-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CCCCN QQPSCXKFDSORFT-IHRRRGAJSA-N 0.000 description 1
- ALEVUGKHINJNIF-QEJZJMRPSA-N Lys-Phe-Ala Chemical compound NCCCC[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](C)C(O)=O)CC1=CC=CC=C1 ALEVUGKHINJNIF-QEJZJMRPSA-N 0.000 description 1
- PDIDTSZKKFEDMB-UWVGGRQHSA-N Lys-Pro-Gly Chemical compound [H]N[C@@H](CCCCN)C(=O)N1CCC[C@H]1C(=O)NCC(O)=O PDIDTSZKKFEDMB-UWVGGRQHSA-N 0.000 description 1
- GHKXHCMRAUYLBS-CIUDSAMLSA-N Lys-Ser-Asn Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(N)=O)C(O)=O GHKXHCMRAUYLBS-CIUDSAMLSA-N 0.000 description 1
- MGKFCQFVPKOWOL-CIUDSAMLSA-N Lys-Ser-Asp Chemical compound C(CCN)C[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(=O)O)C(=O)O)N MGKFCQFVPKOWOL-CIUDSAMLSA-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
- UIJVKVHLCQSPOJ-XIRDDKMYSA-N Lys-Ser-Trp Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](Cc1c[nH]c2ccccc12)C(O)=O UIJVKVHLCQSPOJ-XIRDDKMYSA-N 0.000 description 1
- RMOKGALPSPOYKE-KATARQTJSA-N Lys-Thr-Ser Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CO)C(O)=O RMOKGALPSPOYKE-KATARQTJSA-N 0.000 description 1
- SUZVLFWOCKHWET-CQDKDKBSSA-N Lys-Tyr-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](C)C(O)=O SUZVLFWOCKHWET-CQDKDKBSSA-N 0.000 description 1
- RQILLQOQXLZTCK-KBPBESRZSA-N Lys-Tyr-Gly Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)NCC(O)=O RQILLQOQXLZTCK-KBPBESRZSA-N 0.000 description 1
- MIMXMVDLMDMOJD-BZSNNMDCSA-N Lys-Tyr-Leu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(C)C)C(O)=O MIMXMVDLMDMOJD-BZSNNMDCSA-N 0.000 description 1
- LMMBAXJRYSXCOQ-ACRUOGEOSA-N Lys-Tyr-Phe Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](Cc1ccc(O)cc1)C(=O)N[C@@H](Cc1ccccc1)C(O)=O LMMBAXJRYSXCOQ-ACRUOGEOSA-N 0.000 description 1
- VWPJQIHBBOJWDN-DCAQKATOSA-N Lys-Val-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](C)C(O)=O VWPJQIHBBOJWDN-DCAQKATOSA-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
- TZLYIHDABYBOCJ-FXQIFTODSA-N Met-Asp-Ser Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(O)=O TZLYIHDABYBOCJ-FXQIFTODSA-N 0.000 description 1
- FVKRQMQQFGBXHV-QXEWZRGKSA-N Met-Asp-Val Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O FVKRQMQQFGBXHV-QXEWZRGKSA-N 0.000 description 1
- WRXOPYNEKGZWAZ-FXQIFTODSA-N Met-Ser-Cys Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CS)C(O)=O WRXOPYNEKGZWAZ-FXQIFTODSA-N 0.000 description 1
- QAVZUKIPOMBLMC-AVGNSLFASA-N Met-Val-Leu Chemical compound CSCC[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CC(C)C QAVZUKIPOMBLMC-AVGNSLFASA-N 0.000 description 1
- 102000006399 Metallochaperones Human genes 0.000 description 1
- 108010044086 Metallochaperones Proteins 0.000 description 1
- 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 1
- NWBJYWHLCVSVIJ-UHFFFAOYSA-N N-benzyladenine Chemical compound N=1C=NC=2NC=NC=2C=1NCC1=CC=CC=C1 NWBJYWHLCVSVIJ-UHFFFAOYSA-N 0.000 description 1
- KZNQNBZMBZJQJO-UHFFFAOYSA-N N-glycyl-L-proline Natural products NCC(=O)N1CCCC1C(O)=O KZNQNBZMBZJQJO-UHFFFAOYSA-N 0.000 description 1
- 108010079364 N-glycylalanine Proteins 0.000 description 1
- GRYLNZFGIOXLOG-UHFFFAOYSA-N Nitric acid Chemical compound O[N+]([O-])=O GRYLNZFGIOXLOG-UHFFFAOYSA-N 0.000 description 1
- 240000007594 Oryza sativa Species 0.000 description 1
- 235000007164 Oryza sativa Nutrition 0.000 description 1
- 102000003697 P-type ATPases Human genes 0.000 description 1
- 108090000069 P-type ATPases Proteins 0.000 description 1
- ZWJKVFAYPLPCQB-UNQGMJICSA-N Phe-Arg-Thr Chemical compound C[C@@H](O)[C@H](NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@@H](N)Cc1ccccc1)C(O)=O ZWJKVFAYPLPCQB-UNQGMJICSA-N 0.000 description 1
- WIVCOAKLPICYGY-KKUMJFAQSA-N Phe-Asp-Lys Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N[C@@H](CCCCN)C(=O)O)N WIVCOAKLPICYGY-KKUMJFAQSA-N 0.000 description 1
- KAGCQPSEVAETCA-JYJNAYRXSA-N Phe-Gln-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CC1=CC=CC=C1)N KAGCQPSEVAETCA-JYJNAYRXSA-N 0.000 description 1
- MGBRZXXGQBAULP-DRZSPHRISA-N Phe-Glu-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CCC(O)=O)NC(=O)[C@@H](N)CC1=CC=CC=C1 MGBRZXXGQBAULP-DRZSPHRISA-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
- WFHRXJOZEXUKLV-IRXDYDNUSA-N Phe-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=CC=C1 WFHRXJOZEXUKLV-IRXDYDNUSA-N 0.000 description 1
- CMHTUJQZQXFNTQ-OEAJRASXSA-N Phe-Leu-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)[C@H](CC(C)C)NC(=O)[C@H](CC1=CC=CC=C1)N)O CMHTUJQZQXFNTQ-OEAJRASXSA-N 0.000 description 1
- SCKXGHWQPPURGT-KKUMJFAQSA-N Phe-Lys-Ser Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CO)C(O)=O SCKXGHWQPPURGT-KKUMJFAQSA-N 0.000 description 1
- YVIVIQWMNCWUFS-UFYCRDLUSA-N Phe-Met-Tyr Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)O)NC(=O)[C@H](CC2=CC=CC=C2)N YVIVIQWMNCWUFS-UFYCRDLUSA-N 0.000 description 1
- BPIFSOUEUYDJRM-DCPHZVHLSA-N Phe-Trp-Ala Chemical compound C([C@H](N)C(=O)N[C@@H](CC=1C2=CC=CC=C2NC=1)C(=O)N[C@@H](C)C(O)=O)C1=CC=CC=C1 BPIFSOUEUYDJRM-DCPHZVHLSA-N 0.000 description 1
- 241001503951 Phoma Species 0.000 description 1
- SFECXGVELZFBFJ-VEVYYDQMSA-N Pro-Asp-Thr Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O SFECXGVELZFBFJ-VEVYYDQMSA-N 0.000 description 1
- FKKHDBFNOLCYQM-FXQIFTODSA-N Pro-Cys-Ala Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CS)C(=O)N[C@@H](C)C(O)=O FKKHDBFNOLCYQM-FXQIFTODSA-N 0.000 description 1
- CLNJSLSHKJECME-BQBZGAKWSA-N Pro-Gly-Ala Chemical compound OC(=O)[C@H](C)NC(=O)CNC(=O)[C@@H]1CCCN1 CLNJSLSHKJECME-BQBZGAKWSA-N 0.000 description 1
- MKGIILKDUGDRRO-FXQIFTODSA-N Pro-Ser-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@@H]1CCCN1 MKGIILKDUGDRRO-FXQIFTODSA-N 0.000 description 1
- JXVXYRZQIUPYSA-NHCYSSNCSA-N Pro-Val-Gln Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CCC(N)=O)C(O)=O JXVXYRZQIUPYSA-NHCYSSNCSA-N 0.000 description 1
- MTMJNKFZDQEVSY-BZSNNMDCSA-N Pro-Val-Trp Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(O)=O MTMJNKFZDQEVSY-BZSNNMDCSA-N 0.000 description 1
- 238000002123 RNA extraction Methods 0.000 description 1
- 102000006382 Ribonucleases Human genes 0.000 description 1
- 108010083644 Ribonucleases Proteins 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
- WDXYVIIVDIDOSX-DCAQKATOSA-N Ser-Arg-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)CO)CCCN=C(N)N WDXYVIIVDIDOSX-DCAQKATOSA-N 0.000 description 1
- YPUSXTWURJANKF-KBIXCLLPSA-N Ser-Gln-Ile Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O YPUSXTWURJANKF-KBIXCLLPSA-N 0.000 description 1
- HVKMTOIAYDOJPL-NRPADANISA-N Ser-Gln-Val Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(O)=O HVKMTOIAYDOJPL-NRPADANISA-N 0.000 description 1
- UIGMAMGZOJVTDN-WHFBIAKZSA-N Ser-Gly-Ser Chemical compound OC[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O UIGMAMGZOJVTDN-WHFBIAKZSA-N 0.000 description 1
- 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
- ZOPISOXXPQNOCO-SVSWQMSJSA-N Ser-Ile-Thr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)O)NC(=O)[C@H](CO)N ZOPISOXXPQNOCO-SVSWQMSJSA-N 0.000 description 1
- VMLONWHIORGALA-SRVKXCTJSA-N Ser-Leu-Leu Chemical compound CC(C)C[C@@H](C([O-])=O)NC(=O)[C@H](CC(C)C)NC(=O)[C@@H]([NH3+])CO VMLONWHIORGALA-SRVKXCTJSA-N 0.000 description 1
- GZSZPKSBVAOGIE-CIUDSAMLSA-N Ser-Lys-Ala Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O GZSZPKSBVAOGIE-CIUDSAMLSA-N 0.000 description 1
- LRWBCWGEUCKDTN-BJDJZHNGSA-N Ser-Lys-Ile Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O LRWBCWGEUCKDTN-BJDJZHNGSA-N 0.000 description 1
- AMRRYKHCILPAKD-FXQIFTODSA-N Ser-Met-Asn Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CO)N AMRRYKHCILPAKD-FXQIFTODSA-N 0.000 description 1
- XNXRTQZTFVMJIJ-DCAQKATOSA-N Ser-Met-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CC(C)C)C(O)=O XNXRTQZTFVMJIJ-DCAQKATOSA-N 0.000 description 1
- ZSLFCBHEINFXRS-LPEHRKFASA-N Ser-Met-Pro Chemical compound CSCC[C@@H](C(=O)N1CCC[C@@H]1C(=O)O)NC(=O)[C@H](CO)N ZSLFCBHEINFXRS-LPEHRKFASA-N 0.000 description 1
- ASGYVPAVFNDZMA-GUBZILKMSA-N Ser-Met-Val Chemical compound CC(C)[C@@H](C(=O)O)NC(=O)[C@H](CCSC)NC(=O)[C@H](CO)N ASGYVPAVFNDZMA-GUBZILKMSA-N 0.000 description 1
- AZWNCEBQZXELEZ-FXQIFTODSA-N Ser-Pro-Ser Chemical compound OC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O AZWNCEBQZXELEZ-FXQIFTODSA-N 0.000 description 1
- BMKNXTJLHFIAAH-CIUDSAMLSA-N Ser-Ser-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O BMKNXTJLHFIAAH-CIUDSAMLSA-N 0.000 description 1
- RXUOAOOZIWABBW-XGEHTFHBSA-N Ser-Thr-Arg Chemical compound OC[C@H](N)C(=O)N[C@@H]([C@H](O)C)C(=O)N[C@H](C(O)=O)CCCN=C(N)N RXUOAOOZIWABBW-XGEHTFHBSA-N 0.000 description 1
- PURRNJBBXDDWLX-ZDLURKLDSA-N Ser-Thr-Gly Chemical compound C[C@H]([C@@H](C(=O)NCC(=O)O)NC(=O)[C@H](CO)N)O PURRNJBBXDDWLX-ZDLURKLDSA-N 0.000 description 1
- VLMIUSLQONKLDV-HEIBUPTGSA-N Ser-Thr-Thr Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O VLMIUSLQONKLDV-HEIBUPTGSA-N 0.000 description 1
- OJFFAQFRCVPHNN-JYBASQMISA-N Ser-Thr-Trp Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(O)=O OJFFAQFRCVPHNN-JYBASQMISA-N 0.000 description 1
- JGUWRQWULDWNCM-FXQIFTODSA-N Ser-Val-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CO)C(O)=O JGUWRQWULDWNCM-FXQIFTODSA-N 0.000 description 1
- 241000187747 Streptomyces Species 0.000 description 1
- 241001414851 Susana Species 0.000 description 1
- VFEHSAJCWWHDBH-RHYQMDGZSA-N Thr-Arg-Leu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(C)C)C(O)=O VFEHSAJCWWHDBH-RHYQMDGZSA-N 0.000 description 1
- GKMYGVQDGVYCPC-IUKAMOBKSA-N Thr-Asp-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H]([C@@H](C)O)N GKMYGVQDGVYCPC-IUKAMOBKSA-N 0.000 description 1
- ZUUDNCOCILSYAM-KKHAAJSZSA-N Thr-Asp-Val Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O ZUUDNCOCILSYAM-KKHAAJSZSA-N 0.000 description 1
- HJOSVGCWOTYJFG-WDCWCFNPSA-N Thr-Glu-Lys Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CCCCN)C(=O)O)N)O HJOSVGCWOTYJFG-WDCWCFNPSA-N 0.000 description 1
- AQAMPXBRJJWPNI-JHEQGTHGSA-N Thr-Gly-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)NCC(=O)N[C@@H](CCC(O)=O)C(O)=O AQAMPXBRJJWPNI-JHEQGTHGSA-N 0.000 description 1
- DJDSEDOKJTZBAR-ZDLURKLDSA-N Thr-Gly-Ser Chemical compound C[C@@H](O)[C@H](N)C(=O)NCC(=O)N[C@@H](CO)C(O)=O DJDSEDOKJTZBAR-ZDLURKLDSA-N 0.000 description 1
- KBBRNEDOYWMIJP-KYNKHSRBSA-N Thr-Gly-Thr Chemical compound C[C@H]([C@@H](C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(=O)O)N)O KBBRNEDOYWMIJP-KYNKHSRBSA-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
- XTCNBOBTROGWMW-RWRJDSDZSA-N Thr-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@@H](C)O)N XTCNBOBTROGWMW-RWRJDSDZSA-N 0.000 description 1
- BIBYEFRASCNLAA-CDMKHQONSA-N Thr-Phe-Gly Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@H](C(=O)NCC(O)=O)CC1=CC=CC=C1 BIBYEFRASCNLAA-CDMKHQONSA-N 0.000 description 1
- YGCDFAJJCRVQKU-RCWTZXSCSA-N Thr-Pro-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)[C@@H](C)O YGCDFAJJCRVQKU-RCWTZXSCSA-N 0.000 description 1
- NJGMALCNYAMYCB-JRQIVUDYSA-N Thr-Tyr-Asn Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(N)=O)C(O)=O NJGMALCNYAMYCB-JRQIVUDYSA-N 0.000 description 1
- OGXQLUCMJZSJPW-LYSGOOTNSA-N Trp-Gly-Thr Chemical compound C[C@H]([C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CC1=CNC2=CC=CC=C21)N)O OGXQLUCMJZSJPW-LYSGOOTNSA-N 0.000 description 1
- NRFTYDWKWGJLAR-MELADBBJSA-N Tyr-Asp-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC(=O)O)NC(=O)[C@H](CC2=CC=C(C=C2)O)N)C(=O)O NRFTYDWKWGJLAR-MELADBBJSA-N 0.000 description 1
- NZFCWALTLNFHHC-JYJNAYRXSA-N Tyr-Glu-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@H](CCC(O)=O)NC(=O)[C@@H](N)CC1=CC=C(O)C=C1 NZFCWALTLNFHHC-JYJNAYRXSA-N 0.000 description 1
- JAGGEZACYAAMIL-CQDKDKBSSA-N Tyr-Lys-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](CC1=CC=C(C=C1)O)N JAGGEZACYAAMIL-CQDKDKBSSA-N 0.000 description 1
- PHKQVWWHRYUCJL-HJOGWXRNSA-N Tyr-Phe-Tyr 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](CC1=CC=C(O)C=C1)C(O)=O PHKQVWWHRYUCJL-HJOGWXRNSA-N 0.000 description 1
- QKXAEWMHAAVVGS-KKUMJFAQSA-N Tyr-Pro-Glu Chemical compound N[C@@H](Cc1ccc(O)cc1)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCC(O)=O)C(O)=O QKXAEWMHAAVVGS-KKUMJFAQSA-N 0.000 description 1
- MNWINJDPGBNOED-ULQDDVLXSA-N Tyr-Pro-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)CC1=CC=C(O)C=C1 MNWINJDPGBNOED-ULQDDVLXSA-N 0.000 description 1
- YYLHVUCSTXXKBS-IHRRRGAJSA-N Tyr-Pro-Ser Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O YYLHVUCSTXXKBS-IHRRRGAJSA-N 0.000 description 1
- ITDWWLTTWRRLCC-KJEVXHAQSA-N Tyr-Thr-Arg Chemical compound NC(N)=NCCC[C@@H](C(O)=O)NC(=O)[C@H]([C@H](O)C)NC(=O)[C@@H](N)CC1=CC=C(O)C=C1 ITDWWLTTWRRLCC-KJEVXHAQSA-N 0.000 description 1
- AEOFMCAKYIQQFY-YDHLFZDLSA-N Tyr-Val-Asn Chemical compound NC(=O)C[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CC1=CC=C(O)C=C1 AEOFMCAKYIQQFY-YDHLFZDLSA-N 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
- SMKXLHVZIFKQRB-GUBZILKMSA-N Val-Ala-Met Chemical compound C[C@@H](C(=O)N[C@@H](CCSC)C(=O)O)NC(=O)[C@H](C(C)C)N SMKXLHVZIFKQRB-GUBZILKMSA-N 0.000 description 1
- ZMDCGGKHRKNWKD-LAEOZQHASA-N Val-Asn-Glu Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CCC(=O)O)C(=O)O)N ZMDCGGKHRKNWKD-LAEOZQHASA-N 0.000 description 1
- JLFKWDAZBRYCGX-ZKWXMUAHSA-N Val-Asn-Ser Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)N[C@@H](CO)C(=O)O)N JLFKWDAZBRYCGX-ZKWXMUAHSA-N 0.000 description 1
- HIZMLPKDJAXDRG-FXQIFTODSA-N Val-Cys-Ser Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CO)C(=O)O)N HIZMLPKDJAXDRG-FXQIFTODSA-N 0.000 description 1
- DJEVQCWNMQOABE-RCOVLWMOSA-N Val-Gly-Asp Chemical compound CC(C)[C@@H](C(=O)NCC(=O)N[C@@H](CC(=O)O)C(=O)O)N DJEVQCWNMQOABE-RCOVLWMOSA-N 0.000 description 1
- MDYSKHBSPXUOPV-JSGCOSHPSA-N Val-Gly-Phe Chemical compound CC(C)[C@@H](C(=O)NCC(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)N MDYSKHBSPXUOPV-JSGCOSHPSA-N 0.000 description 1
- KZKMBGXCNLPYKD-YEPSODPASA-N Val-Gly-Thr Chemical compound CC(C)[C@H](N)C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(O)=O KZKMBGXCNLPYKD-YEPSODPASA-N 0.000 description 1
- PTFPUAXGIKTVNN-ONGXEEELSA-N Val-His-Gly Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)NCC(=O)O)N PTFPUAXGIKTVNN-ONGXEEELSA-N 0.000 description 1
- HQYVQDRYODWONX-DCAQKATOSA-N Val-His-Ser Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)N[C@@H](CO)C(=O)O)N HQYVQDRYODWONX-DCAQKATOSA-N 0.000 description 1
- VHRLUTIMTDOVCG-PEDHHIEDSA-N Val-Ile-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)CC)C(=O)O)NC(=O)[C@H](C(C)C)N VHRLUTIMTDOVCG-PEDHHIEDSA-N 0.000 description 1
- OVBMCNDKCWAXMZ-NAKRPEOUSA-N Val-Ile-Ser Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](C(C)C)N OVBMCNDKCWAXMZ-NAKRPEOUSA-N 0.000 description 1
- APQIVBCUIUDSMB-OSUNSFLBSA-N Val-Ile-Thr Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)O)NC(=O)[C@H](C(C)C)N APQIVBCUIUDSMB-OSUNSFLBSA-N 0.000 description 1
- MYLNLEIZWHVENT-VKOGCVSHSA-N Val-Ile-Trp Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)O)NC(=O)[C@H](C(C)C)N MYLNLEIZWHVENT-VKOGCVSHSA-N 0.000 description 1
- OTJMMKPMLUNTQT-AVGNSLFASA-N Val-Leu-Arg Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)NC(=O)[C@H](C(C)C)N OTJMMKPMLUNTQT-AVGNSLFASA-N 0.000 description 1
- FEXILLGKGGTLRI-NHCYSSNCSA-N Val-Leu-Asn Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](C(C)C)N FEXILLGKGGTLRI-NHCYSSNCSA-N 0.000 description 1
- BTWMICVCQLKKNR-DCAQKATOSA-N Val-Leu-Ser Chemical compound CC(C)[C@H]([NH3+])C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C([O-])=O BTWMICVCQLKKNR-DCAQKATOSA-N 0.000 description 1
- RWOGENDAOGMHLX-DCAQKATOSA-N Val-Lys-Ala Chemical compound C[C@@H](C(=O)O)NC(=O)[C@H](CCCCN)NC(=O)[C@H](C(C)C)N RWOGENDAOGMHLX-DCAQKATOSA-N 0.000 description 1
- MJFSRZZJQWZHFQ-SRVKXCTJSA-N Val-Met-Val Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCSC)C(=O)N[C@@H](C(C)C)C(=O)O)N MJFSRZZJQWZHFQ-SRVKXCTJSA-N 0.000 description 1
- NHXZRXLFOBFMDM-AVGNSLFASA-N Val-Pro-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)C(C)C NHXZRXLFOBFMDM-AVGNSLFASA-N 0.000 description 1
- QZKVWWIUSQGWMY-IHRRRGAJSA-N Val-Ser-Phe Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 QZKVWWIUSQGWMY-IHRRRGAJSA-N 0.000 description 1
- HWNYVQMOLCYHEA-IHRRRGAJSA-N Val-Ser-Tyr Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC1=CC=C(C=C1)O)C(=O)O)N HWNYVQMOLCYHEA-IHRRRGAJSA-N 0.000 description 1
- NZYNRRGJJVSSTJ-GUBZILKMSA-N Val-Ser-Val Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(O)=O NZYNRRGJJVSSTJ-GUBZILKMSA-N 0.000 description 1
- IECQJCJNPJVUSB-IHRRRGAJSA-N Val-Tyr-Ser Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](Cc1ccc(O)cc1)C(=O)N[C@@H](CO)C(O)=O IECQJCJNPJVUSB-IHRRRGAJSA-N 0.000 description 1
- DFQZDQPLWBSFEJ-LSJOCFKGSA-N Val-Val-Asn Chemical compound CC(C)[C@@H](C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(=O)N)C(=O)O)N DFQZDQPLWBSFEJ-LSJOCFKGSA-N 0.000 description 1
- ZLNYBMWGPOKSLW-LSJOCFKGSA-N Val-Val-Asp Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(O)=O)C(O)=O ZLNYBMWGPOKSLW-LSJOCFKGSA-N 0.000 description 1
- 238000002835 absorbance Methods 0.000 description 1
- 108010024078 alanyl-glycyl-serine Proteins 0.000 description 1
- 108010005233 alanylglutamic acid Proteins 0.000 description 1
- 108010087924 alanylproline Proteins 0.000 description 1
- 108010070783 alanyltyrosine Proteins 0.000 description 1
- 150000001413 amino acids Chemical class 0.000 description 1
- 230000003698 anagen phase Effects 0.000 description 1
- 238000004458 analytical method Methods 0.000 description 1
- 238000013459 approach Methods 0.000 description 1
- 108010062796 arginyllysine Proteins 0.000 description 1
- 108010060035 arginylproline Proteins 0.000 description 1
- 108010077245 asparaginyl-proline Proteins 0.000 description 1
- 108010038633 aspartylglutamate Proteins 0.000 description 1
- 108010047857 aspartylglycine Proteins 0.000 description 1
- 230000009286 beneficial effect Effects 0.000 description 1
- 238000009395 breeding Methods 0.000 description 1
- 230000001488 breeding effect Effects 0.000 description 1
- 238000010804 cDNA synthesis Methods 0.000 description 1
- 239000001506 calcium phosphate Substances 0.000 description 1
- 229910000389 calcium phosphate Inorganic materials 0.000 description 1
- 235000011010 calcium phosphates Nutrition 0.000 description 1
- 238000004113 cell culture Methods 0.000 description 1
- 235000013339 cereals Nutrition 0.000 description 1
- 239000003153 chemical reaction reagent Substances 0.000 description 1
- 238000003776 cleavage reaction Methods 0.000 description 1
- 238000000975 co-precipitation Methods 0.000 description 1
- 238000010276 construction Methods 0.000 description 1
- 238000005520 cutting process Methods 0.000 description 1
- 108010060199 cysteinylproline Proteins 0.000 description 1
- 239000008367 deionised water Substances 0.000 description 1
- 229910021641 deionized water Inorganic materials 0.000 description 1
- 238000012217 deletion Methods 0.000 description 1
- 230000037430 deletion Effects 0.000 description 1
- 238000001514 detection method Methods 0.000 description 1
- 238000011161 development Methods 0.000 description 1
- 108010054812 diprotin A Proteins 0.000 description 1
- 108010054813 diprotin B Proteins 0.000 description 1
- 238000005538 encapsulation Methods 0.000 description 1
- 101150100366 end gene Proteins 0.000 description 1
- 238000000605 extraction Methods 0.000 description 1
- 229910001447 ferric ion Inorganic materials 0.000 description 1
- 238000007667 floating Methods 0.000 description 1
- 239000006260 foam Substances 0.000 description 1
- 229940105450 folic acid 1 mg/ml Drugs 0.000 description 1
- 235000013305 food Nutrition 0.000 description 1
- 238000010230 functional analysis Methods 0.000 description 1
- 230000002538 fungal effect Effects 0.000 description 1
- 108010006664 gamma-glutamyl-glycyl-glycine Proteins 0.000 description 1
- 238000003209 gene knockout Methods 0.000 description 1
- 238000010353 genetic engineering Methods 0.000 description 1
- 108010049041 glutamylalanine Proteins 0.000 description 1
- 108010089804 glycyl-threonine Proteins 0.000 description 1
- 108010010147 glycylglutamine Proteins 0.000 description 1
- 108010077515 glycylproline Proteins 0.000 description 1
- 239000001963 growth medium Substances 0.000 description 1
- 230000036541 health Effects 0.000 description 1
- 230000003301 hydrolyzing effect Effects 0.000 description 1
- 238000000338 in vitro Methods 0.000 description 1
- 238000001727 in vivo Methods 0.000 description 1
- 238000011534 incubation Methods 0.000 description 1
- 230000006698 induction Effects 0.000 description 1
- 238000009616 inductively coupled plasma Methods 0.000 description 1
- 230000003993 interaction Effects 0.000 description 1
- 150000002500 ions Chemical class 0.000 description 1
- 108010078274 isoleucylvaline Proteins 0.000 description 1
- 108010077158 leucinyl-arginyl-tryptophan Proteins 0.000 description 1
- 108010000761 leucylarginine Proteins 0.000 description 1
- 230000000670 limiting effect Effects 0.000 description 1
- 239000002502 liposome Substances 0.000 description 1
- 108010009298 lysylglutamic acid Proteins 0.000 description 1
- 108010017391 lysylvaline Proteins 0.000 description 1
- 210000004962 mammalian cell Anatomy 0.000 description 1
- 210000001161 mammalian embryo Anatomy 0.000 description 1
- 239000003550 marker Substances 0.000 description 1
- 238000005259 measurement Methods 0.000 description 1
- 238000010297 mechanical methods and process Methods 0.000 description 1
- 230000007246 mechanism Effects 0.000 description 1
- 108020004999 messenger RNA Proteins 0.000 description 1
- 229910052751 metal Inorganic materials 0.000 description 1
- 239000002184 metal Substances 0.000 description 1
- 108010005942 methionylglycine Proteins 0.000 description 1
- 238000000520 microinjection Methods 0.000 description 1
- 238000002156 mixing Methods 0.000 description 1
- 238000010369 molecular cloning Methods 0.000 description 1
- 230000009456 molecular mechanism Effects 0.000 description 1
- 229910017604 nitric acid Inorganic materials 0.000 description 1
- 210000000056 organ Anatomy 0.000 description 1
- 230000000737 periodic effect Effects 0.000 description 1
- 108010084572 phenylalanyl-valine Proteins 0.000 description 1
- 108010012581 phenylalanylglutamate Proteins 0.000 description 1
- 108010073101 phenylalanylleucine Proteins 0.000 description 1
- 108010051242 phenylalanylserine Proteins 0.000 description 1
- 230000029553 photosynthesis Effects 0.000 description 1
- 238000010672 photosynthesis Methods 0.000 description 1
- 108010025488 pinealon Proteins 0.000 description 1
- 230000008121 plant development Effects 0.000 description 1
- 238000003752 polymerase chain reaction Methods 0.000 description 1
- 229920001184 polypeptide Polymers 0.000 description 1
- 238000002360 preparation method Methods 0.000 description 1
- 238000004321 preservation Methods 0.000 description 1
- 108090000765 processed proteins & peptides Proteins 0.000 description 1
- 102000004196 processed proteins & peptides Human genes 0.000 description 1
- 210000001236 prokaryotic cell Anatomy 0.000 description 1
- 108010031719 prolyl-serine Proteins 0.000 description 1
- 108010029020 prolylglycine Proteins 0.000 description 1
- 238000010188 recombinant method Methods 0.000 description 1
- 230000008844 regulatory mechanism Effects 0.000 description 1
- 230000010076 replication Effects 0.000 description 1
- 238000010839 reverse transcription Methods 0.000 description 1
- 235000009566 rice Nutrition 0.000 description 1
- 239000012883 rooting culture medium Substances 0.000 description 1
- 239000012882 rooting medium Substances 0.000 description 1
- 230000007017 scission Effects 0.000 description 1
- 108010048397 seryl-lysyl-leucine Proteins 0.000 description 1
- 108010071207 serylmethionine Proteins 0.000 description 1
- 238000009331 sowing Methods 0.000 description 1
- 238000001228 spectrum Methods 0.000 description 1
- 238000004659 sterilization and disinfection Methods 0.000 description 1
- 238000003860 storage Methods 0.000 description 1
- 238000013518 transcription Methods 0.000 description 1
- 230000035897 transcription Effects 0.000 description 1
- 230000002103 transcriptional effect Effects 0.000 description 1
- 238000003151 transfection method Methods 0.000 description 1
- 238000012546 transfer Methods 0.000 description 1
- 230000014621 translational initiation Effects 0.000 description 1
- QORWJWZARLRLPR-UHFFFAOYSA-H tricalcium bis(phosphate) Chemical compound [Ca+2].[Ca+2].[Ca+2].[O-]P([O-])([O-])=O.[O-]P([O-])([O-])=O QORWJWZARLRLPR-UHFFFAOYSA-H 0.000 description 1
- 108010080629 tryptophan-leucine Proteins 0.000 description 1
- 108010038745 tryptophylglycine Proteins 0.000 description 1
- 108010051110 tyrosyl-lysine Proteins 0.000 description 1
- 210000003934 vacuole Anatomy 0.000 description 1
- 108010073969 valyllysine Proteins 0.000 description 1
- 230000017260 vegetative to reproductive phase transition of meristem Effects 0.000 description 1
- 238000005303 weighing Methods 0.000 description 1
- 210000005253 yeast cell Anatomy 0.000 description 1
Classifications
-
- C—CHEMISTRY; METALLURGY
- C07—ORGANIC CHEMISTRY
- C07K—PEPTIDES
- C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
- C07K14/415—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from plants
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01G—HORTICULTURE; CULTIVATION OF VEGETABLES, FLOWERS, RICE, FRUIT, VINES, HOPS OR SEAWEED; FORESTRY; WATERING
- A01G22/00—Cultivation of specific crops or plants not otherwise provided for
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8262—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield involving plant development
- C12N15/827—Flower development or morphology, e.g. flowering promoting factor [FPF]
-
- C—CHEMISTRY; METALLURGY
- C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
- C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
- C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
- C12N15/09—Recombinant DNA-technology
- C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
- C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
- C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
- C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
- C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
- C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
-
- Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
- Y02—TECHNOLOGIES OR APPLICATIONS FOR MITIGATION OR ADAPTATION AGAINST CLIMATE CHANGE
- Y02A—TECHNOLOGIES FOR ADAPTATION TO CLIMATE CHANGE
- Y02A40/00—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production
- Y02A40/10—Adaptation technologies in agriculture, forestry, livestock or agroalimentary production in agriculture
- Y02A40/146—Genetically Modified [GMO] plants, e.g. transgenic plants
Landscapes
- Health & Medical Sciences (AREA)
- Life Sciences & Earth Sciences (AREA)
- Genetics & Genomics (AREA)
- Chemical & Material Sciences (AREA)
- Organic Chemistry (AREA)
- Engineering & Computer Science (AREA)
- Molecular Biology (AREA)
- General Engineering & Computer Science (AREA)
- Bioinformatics & Cheminformatics (AREA)
- Biotechnology (AREA)
- Wood Science & Technology (AREA)
- Biomedical Technology (AREA)
- Zoology (AREA)
- General Health & Medical Sciences (AREA)
- Biochemistry (AREA)
- Biophysics (AREA)
- Physics & Mathematics (AREA)
- Botany (AREA)
- Cell Biology (AREA)
- Microbiology (AREA)
- Plant Pathology (AREA)
- Proteomics, Peptides & Aminoacids (AREA)
- Medicinal Chemistry (AREA)
- Gastroenterology & Hepatology (AREA)
- Physiology (AREA)
- Environmental Sciences (AREA)
- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
- Micro-Organisms Or Cultivation Processes Thereof (AREA)
Abstract
本发明公开了一种重金属转运蛋白PhHMA5II‑1及其相关产品和用途。所述蛋白为如下(1)或(2)的蛋白：(1)由如SEQ ID NO.2所示的氨基酸序列组成的蛋白；(2)将如SEQ ID NO.2所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由(1)衍生的蛋白。本发明的重金属转运蛋白PhHMA5II‑1的靶序列，经基因编辑技术与pYLCRISPR/Cas9载体连接后导入矮牵牛中形成矮牵牛突变体，经铜胁迫后，该矮牵牛突变体的花瓣中的铁含量明显降低，铜和锰含量也有不同程度的下降；同时矮牵牛突变体的生长得到了促进，特别是花瓣的数量性状得到了提高，尤其是花瓣的重量。
Description
技术领域
本发明涉及基因工程技术领域，特别是涉及一种重金属转运蛋白PhHMA5II-1及其相关产品和用途。
背景技术
重金属如铜，是我国土壤的重要污染源之一，过量的铜会严重影响植物的生长。若铜通过食物链进入人体内，将影响人体的健康。所以在铜污染严重的土壤上，用花卉等经济作物代替粮食作物种植能够有效合理的利用土地。
高浓度铜，能够引起植物体叶片和花瓣内铁离子平衡的失调，从而导致植物的光合作用速率降低，生物量降低，从而严重影响植物的生长和发育。但由于对植物特别是花瓣内中重金属，例如铜转运的分子机理，特别是铁和铜离子互作的分子调控机理尚不清楚，严重影响了抗性花卉植物的筛选和选育。
矮牵牛是重要的园林造景植物，因其花色丰富，花期长等特点，是世界最受欢迎的道路和园林布景花卉，有花坛之王的美誉。HMA5属于植物重金属转运家族蛋白，是植物中最重要的铜解毒蛋白之一。它能够利用水解ATP产生的能量将单价的铜离子泵出细胞或者泵入液泡，从而达到铜解毒的作用。但目前对其功能机理的研究主要集中在拟南芥和水稻等植物的根部。然而，其在花卉植物中的研究尚少，尤其是在花瓣中的提高重金属转运能力或提高重金属耐受性的研究很少。此外，HMA5家族蛋白转运铁的功能也未见报道。
发明内容
鉴于以上所述现有技术的缺点，本发明的目的在于提供一种重金属转运蛋白PhHMA5II-1及其相关产品和用途，进而解决现有技术中的问题。
本发明目的之一在于提供一种重金属转运蛋白PhHMA5II-1，所述蛋白为如下(1)或(2)的蛋白：
(1)由如SEQ ID NO.2所示的氨基酸序列组成的蛋白；
(2)将如SEQ ID NO.2所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由(1)衍生的蛋白。
本发明目的之二在于提供上述所述蛋白相关的生物材料，包括如下任一种：
a)编码如上述所述蛋白的多核苷酸；
b)含有a)所述多核苷酸的重组表达载体；
c)含有a)所述多核苷酸的生物工程菌，或含有b)所述重组表达载体的生物工程菌；
d)含有a)所述多核苷酸的转基因植物细胞，或含有b)所述重组表达载体的转基因植物。
根据本申请的技术方案，a)中，所述多核苷酸包括如SEQ ID NO.1所示序列。
本发明目的之三在于提供如上述所述蛋白作为靶标用于提高植物对重金属耐受性或提高植物重金属转运能力中的用途；优选包括促进植物生长，包括促进根系生长、提高花瓣的数量性状。本申请中，花瓣的数量性状包括花径、花型、花瓣大小和花瓣重量。
本发明目的之四在于提供如上述所述的蛋白质或如上述所述的生物材料在筛选用于提高植物对重金属耐受性或提高植物重金属转运能力产品中的用途；优选包括促进植物生长，包括促进根系生长、提高花朵的数量性状。
本发明目的之五在于提供一种用于提高植物对重金属耐受性或提高植物重金属转运能力的产品，包括PhHMA5II-1抑制剂或PhHMA5II-1基因干扰核酸构建体；
所述PhHMA5II-1抑制剂是指使权利要求1所述的蛋白PhHMA5II-1质突变或失活或功能下降的化合物；
或，所述PhHMA5II-1基因干扰核酸构建体是指使权利要求1所述的蛋白PhHMA5II-1质突变或失活或功能下降的编辑系统。
本发明目的之六在于提供一种构建转基因植物的方法，向植物导入上述所述的PhHMA5II-1抑制剂或PhHMA5II-1基因干扰核酸构建体，得到植物突变体。
根据本申请的技术方案，采用农杆菌介导法将如上述所述的PhHMA5II-1基因干扰核酸构建体导入所述植物。
优选地，所述农杆菌介导法为：
将上述所述的PhHMA5II-1基因干扰核酸构建体转化于农杆菌；
将含有所述PhHMA5II-1基因干扰核酸构建体的农杆菌导入所述植物。
本发明目的之七在于提供一种培育对重金属耐受性高或对重金属转运能力高的植物的方法，在铜胁迫下，种植如上述所述的植物突变体，进行培育。
根据本申请的技术方案，所述铜选自硫酸铜。具体地，所述硫酸铜的浓度不高于10μM。
根据本申请的技术方案，所述植物包括双子叶植物或单子叶植物。
在一个具体实施方式中，所述植物为双子叶植物。
较佳的，所述双子叶植物选自拟南芥、紫花苜宿、矮牵牛、棉花、蓖麻和矮牵牛。具体地，所述双子叶植物为矮牵牛。
根据本申请的技术方案，所述重金属包括铜、锌、锰和铁。
在一个具体实施方式中，所述重金属为铁和铜。
较佳的，所述重金属为铁。
本发明具有以下有益效果：
本发明从矮牵牛W115中获得一个重金属转运基因PhHMA5II-1，该基因与拟南芥AtHMA5基因具有较高的同源性。该基因的靶序列，经基因编辑技术与pYLCRISPR/Cas9载体连接后导入矮牵牛中形成矮牵牛突变体，在高铜胁迫下，该矮牵牛突变体的花瓣中的铁含量明显降低，铜和锰含量也有不同程度的下降；同时该矮牵牛突变体的生长得到了促进，特别是花瓣的数量性状得到了提高，尤其是花瓣的重量。
附图说明
图1显示为本发明实施例1的矮牵牛PhHMA5II-1氨基酸序列与拟南芥AtHMA5氨基酸序列的比较图。
图2显示为本发明实施例1的矮牵牛PhHMA5II-1蛋白的功能结构域图。
图3显示为本发明实施例2的重组质粒的谱图。
图4显示为本发明实施例3的矮牵牛突变体和野生型矮牵牛的部分基因序列对比图。
图5显示为本发明实施例4中在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣中铜离子、锰离子和锌离子的含量图。
图6显示为本发明实施例4中在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣中铁离子的含量图。
图7显示为本发明实施例4中在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣干重图。
具体实施方式
以下通过特定的具体实例说明本发明的实施方式，本领域技术人员可由本说明书所揭露的内容轻易地了解本发明的其他优点与功效。本发明还可以通过另外不同的具体实施方式加以实施或应用，本说明书中的各项细节也可以基于不同观点与应用，在没有背离本发明的精神下进行各种修饰或改变。
在进一步描述本发明具体实施方式之前，应理解，本发明的保护范围不局限于下述特定的具体实施方案；还应当理解，本发明实施例中使用的术语是为了描述特定的具体实施方案，而不是为了限制本发明的保护范围；在本发明说明书和权利要求书中，除非文中另外明确指出，单数形式“一个”、“一”和“这个”包括复数形式。
当实施例给出数值范围时，应理解，除非本发明另有说明，每个数值范围的两个端点以及两个端点之间任何一个数值均可选用。除非另外定义，本发明中使用的所有技术和科学术语与本技术领域技术人员通常理解的意义相同。除实施例中使用的具体方法、设备、材料外，根据本技术领域的技术人员对现有技术的掌握及本发明的记载，还可以使用与本发明实施例中所述的方法、设备、材料相似或等同的现有技术的任何方法、设备和材料来实现本发明。
除非另外说明，本发明中所公开的实验方法、检测方法、制备方法均采用本技术领域常规的分子生物学、生物化学、染色质结构和分析、分析化学、细胞培养、重组DNA技术及相关领域的常规技术。这些技术在现有文献中已有完善说明，具体可参见Sambrook等MOLECULAR CLONING：A LABORATORY MANUAL，Second edition，Cold Spring HarborLaboratory Press，1989and Third edition，2001；Ausubel等，CURRENT PROTOCOLS INMOLECULAR BIOLOGY，John Wiley&Sons，New York，1987and periodic updates；theseries METHODS IN ENZYMOLOGY，Academic Press，San Diego；Wolffe，CHROMATINSTRUCTURE AND FUNCTION，Third edition，Academic Press，San Diego，1998；METHODS INENZYMOLOGY，Vol.304，Chromatin(P.M.Wassarman and A.P.Wolffe，eds.)，AcademicPress，San Diego，1999；和METHODS IN MOLECULAR BIOLOGY，Vol.119，ChromatinProtocols(P.B.Becker，ed.)Humana Press，Totowa，1999等。
本发明实施例的目的在于填补矮牵牛“W115”PhHMA5II-1基因的克隆、以及基因的功能分析的空白。
本发明的第一方面，提供了一种重金属转运蛋白PhHMA5II-1，所述蛋白为如下(1)或(2)的蛋白：(1)由如SEQ ID NO.2所示的氨基酸序列组成的蛋白；(2)将如SEQ ID NO.2所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加且具有相同功能的由(1)衍生的蛋白。
SEQ ID NO：2所示序列具体为：
MATAKFLSFACIRNESKYGDFSSRPHYPSMPKYPKGVTISTDEEKSIHGTESKAIYSVHGMSCSACAGSVEKAIKRLAGIKEAVVDVLNNRAQVIFYPSYVNEEMIRETIEDVGFQATLITEEATEKSSQVCRIRIKGMTCTSCSTTVESALQLIPGVQKAQVALATEEAEIQYDPRILTYNQLLETIENTGFEAILISTGEDRSKILLKIDGVHTGDSMRIIESSLRALPGVEDIDIDPELKKLSVSYKSDIIGPRDFIRVIESTGSGQFNAMLYPEGGGEQSHRQEEIGNYRRSFLWSLFFTIPVFLTSMVFMYIPGLKDGLDIKVVNMLSIGEILRWVLSTPVQFIIGRRFYSGSYKALRHGSANMDVLIALGTNAAYFYSVYSVLRAATSPSFKSTDFFETSSMLISFILLGKYLEVLAKGKTSEAIAKLMNLTPDTATLLQHGDEGNVVNEEEIDSRLIQKNDVIRIVPGAKVACDGFVIWGQSHVNESMITGESRPVAKRKGDIVIGGTVNENGVLHIRATRVGSESALSQIVRLVESAQMAKAPVQKFADGISKYFVPLVIIVSFSTWLAWFLAGKYAAYPKSWIPSSMDSFQLALQFGISVMVIACPCALGLATPTAVMVGTGVGASRGVLIKGGQALESAQKVNCIVFDKTGTLTMGKPVVVNTRLFRTMVLREFYELVAAAEVNSEHPLAKAIVEYAKKFREDEENPVWPEIQDFESITGHGVKAIVHSKKVIVGNKSLMLAQGISVSADADEVLAETEELAQTGILVSVDGVLTGVVAISDPVKPGAREVISLLKSMNVESKLVTGDNWGTANAIAKEVGITDVIAEAKPEDKAEKVKELQSSGKVVAMVGDGINDSPALVAADVGMAIGAGTDIAIEAADIVLMKSNLEDVITAIDLSRKTFGRIRLNYFWAFGYNLLGIPIAAGALFPSTRFRLPPWVAGAAMAASSVSVVCSSLLLKNYTRPKKLDNLGIGGITVE。
本发明第二方面，还提供了与第一方面所述蛋白相关的生物材料，包括如下任一种：
a)编码如第一方面所述蛋白的多核苷酸；
b)含有a)所述多核苷酸的重组表达载体；
c)含有a)所述多核苷酸的生物工程菌，或含有b)所述重组表达载体的生物工程菌；
d)含有a)所述多核苷酸的转基因植物细胞，或含有b)所述重组表达载体的转基因植物。
本发明的多核苷酸可以是DNA形式或RNA形式，DNA形式包括cDNA、基因组DNA或人工合成的DNA。DNA可以是单链的或是双链的。DNA可以是编码链或非编码链。编码成熟多肽的编码区序列可以与SEQ ID NO:1中的编码区序列相同或者是简并的变异体。如本文所用，“简并的变异体”在本发明中是指编码具有SEQ ID NO:2所示序列的蛋白，但与SEQ ID NO:1中的编码区序列有差别的核酸序列。在本发明的一个具体实施例中，所述多核苷酸从矮牵牛“W115”的组织中提取出来。所述多核苷酸具体为PhHMA5II-1基因。
SEQ ID NO：1所示序列具体为：
ATGGCAACTGCAAAGTTTCTTTCATTTGCATGCATAAGAAATGAAAGTAAATATGGAGATTTTTCATCAAGACCACATTATCCATCAATGCCAAAATATCCAAAGGGTGTTACTATATCAACTGATGAAGAAAAAAGTATACATGGAACAGAATCAAAGGCAATATATTCAGTACATGGAATGAGTTGTTCTGCATGTGCAGGTTCTGTAGAGAAAGCAATTAAAAGACTTGCTGGTATTAAAGAAGCTGTTGTTGATGTCTTGAATAATAGAGCTCAAGTTATTTTTTATCCTAGCTATGTTAATGAGGAAATGATCCGTGAGACCATAGAAGATGTTGGATTCCAGGCTACTCTGATTACAGAGGAGGCAACGGAGAAATCCTCCCAAGTCTGCCGGATACGTATAAAAGGAATGACCTGCACTTCTTGCTCCACAACTGTTGAATCTGCTTTACAGTTGATCCCAGGTGTACAGAAAGCACAAGTAGCATTAGCAACTGAAGAGGCTGAAATTCAATATGATCCAAGGATATTAACTTACAATCAGCTTTTAGAGACCATAGAAAATACTGGATTTGAAGCCATATTAATTAGCACAGGAGAAGATAGGAGCAAAATATTGCTGAAAATTGATGGAGTACACACTGGCGATTCCATGAGGATCATTGAAAGTTCTCTTAGAGCACTCCCGGGTGTTGAAGACATTGATATTGATCCTGAGTTGAAGAAGTTATCCGTTTCTTATAAATCAGATATAATAGGACCTAGAGATTTCATTCGAGTAATTGAGTCAACTGGTTCCGGACAGTTCAATGCAATGTTATATCCTGAAGGAGGTGGGGAACAATCGCATAGACAGGAGGAAATTGGAAACTATCGTCGGTCCTTTCTCTGGAGCTTGTTTTTTACCATTCCAGTTTTCTTGACTTCCATGGTCTTTATGTATATTCCTGGTCTGAAGGATGGATTGGATATTAAAGTTGTTAACATGCTTAGCATTGGAGAGATTTTGCGGTGGGTGCTTTCAACTCCAGTGCAATTCATCATTGGCCGTCGTTTCTATTCTGGTTCTTACAAAGCATTACGTCATGGTTCTGCAAATATGGATGTTTTGATTGCTTTGGGAACAAATGCGGCCTACTTTTATTCAGTCTACTCGGTGTTGAGAGCTGCTACTTCCCCAAGCTTCAAGTCTACTGATTTTTTTGAGACCAGCTCAATGCTCATTTCCTTCATTCTACTCGGGAAGTATCTTGAAGTTTTGGCCAAAGGGAAGACATCGGAGGCCATTGCTAAACTCATGAACTTGACCCCTGATACTGCAACATTATTACAGCATGGTGATGAAGGAAATGTGGTAAATGAGGAAGAAATTGATAGCCGACTGATACAGAAGAATGATGTGATCAGAATCGTTCCAGGTGCAAAAGTAGCCTGTGATGGTTTTGTCATCTGGGGACAAAGCCACGTCAACGAGAGTATGATAACCGGAGAATCTCGGCCAGTGGCCAAACGGAAGGGCGACATAGTGATTGGAGGAACTGTGAATGAGAATGGTGTTCTGCATATTAGAGCTACTAGAGTTGGATCAGAGAGTGCGCTTTCACAGATTGTTAGGCTGGTTGAATCAGCACAGATGGCTAAAGCTCCTGTTCAAAAATTTGCTGATGGCATTTCTAAATATTTTGTGCCTCTTGTTATTATTGTCTCCTTTTCCACTTGGCTAGCCTGGTTTTTAGCTGGGAAATACGCCGCCTATCCTAAATCCTGGATTCCATCTTCCATGGATAGCTTTCAGCTTGCCTTGCAATTTGGTATATCTGTTATGGTCATAGCTTGCCCGTGTGCCCTTGGTCTTGCTACTCCAACTGCTGTCATGGTTGGCACAGGAGTTGGTGCTTCTCGTGGTGTCCTGATTAAAGGTGGGCAGGCTTTGGAAAGCGCTCAAAAGGTGAACTGTATAGTGTTTGATAAGACGGGTACACTTACAATGGGGAAACCAGTCGTTGTAAATACAAGGCTTTTCAGAACTATGGTACTTCGAGAATTTTATGAATTGGTTGCTGCAGCTGAGGTAAATAGTGAGCACCCCTTAGCCAAGGCAATTGTCGAATATGCCAAAAAGTTTAGAGAGGATGAGGAGAACCCGGTATGGCCTGAAATCCAGGACTTTGAGTCAATAACTGGCCATGGCGTGAAGGCTATTGTCCACAGCAAGAAAGTAATAGTTGGAAACAAGAGCTTGATGTTAGCACAAGGCATTTCTGTTTCAGCTGATGCCGATGAAGTTCTAGCTGAAACAGAAGAATTGGCTCAAACTGGAATACTTGTATCAGTTGATGGTGTACTGACTGGAGTTGTTGCTATATCAGATCCTGTGAAGCCCGGAGCTCGTGAAGTCATTTCTCTTCTTAAGTCTATGAATGTCGAGAGTAAACTAGTAACAGGTGACAATTGGGGAACAGCTAATGCCATTGCCAAGGAAGTTGGCATTACGGATGTTATTGCAGAAGCAAAACCTGAAGACAAAGCAGAAAAAGTGAAGGAATTGCAGAGTTCAGGAAAAGTTGTTGCAATGGTCGGAGATGGAATTAACGATTCTCCAGCTCTCGTAGCAGCAGATGTTGGAATGGCAATAGGGGCGGGGACAGATATAGCTATTGAGGCAGCTGATATTGTCCTAATGAAAAGCAATCTGGAAGATGTCATAACTGCTATTGACCTTTCCAGGAAAACATTTGGTCGAATTCGTCTGAACTACTTTTGGGCATTTGGCTATAACCTTCTTGGCATCCCAATTGCTGCTGGAGCTCTTTTCCCATCTACTAGATTTCGCCTGCCACCGTGGGTAGCAGGTGCAGCAATGGCTGCTTCTTCAGTGAGTGTTGTCTGCAGCTCTCTTTTGTTGAAGAATTACACGAGACCAAAGAAGCTTGATAATCTTGGGATTGGAGGCATAACTGTTGAGTGA。
本发明对矮牵牛基因进行分析，获得一个转运重金属的基因，具体为PhHMA5II-1基因。PhHMA5II-1基因与拟南芥AtHMA5的基因相似性极高。
在本发明中，矮牵牛“W115”的PhHMA5II-1基因的多核苷酸全长序列或其片段可以用PCR扩增法、重组法或人工合成的方法获得。对于PCR扩增法，可根据本发明的实施例记载的有关核苷酸序列，尤其是开放阅读框序列来设计引物，并用市售的cDNA库或按本领域技术人员已知的常规方法所制备的cDNA作为模板，扩增而得有关序列。当序列较长时，常常需要进行两次或多次PCR扩增，然后再将各次扩增出的片段按正确次序拼接在一起。
此外，可以根据本发明实施例记载的矮牵牛“W115”的PhHMA5II-1核苷酸序列，在核酸同源性的基础上，筛选矮牵牛“W115”的PhHMA5II-1相关同源基因。
在本发明一个具体实施例中，所述SEQ ID NO：1所示的多核苷酸通过包括如下步骤的方法获得：以矮牵牛的cDNA为模板进行PCR扩增，所述PCR扩增的正向引物如序列SEQID NO：8所示，所述PCR扩增的反向引物如序列SEQ ID NO：9所示。
SEQ ID NO：8所示序列具体为：
5'-GGGGACAAGTTTGTACAAAAAAGCAGGCTCAATGGCAACTGCAAAGTTTCTTTC-3'
SEQ ID NO：9所示序列具体为：
5'-GGGGACCACTTTGTACAAGAAAGCTGGGTTCACTCAACAGTTATGCCTCCAATC-3'
需要说明的是，在本发明实施例中，“多核苷酸”可以是指，该多核苷酸已从天然状态下位于其两侧的序列中分离出来，也可以指该多核苷酸已经与天然状态下伴随核酸的组分分开，而且已经与在细胞中相伴随的蛋白质分开。
本发明的含有a)所述多核苷酸的重组表达载体。本发明中，编码所述蛋白质的多核苷酸序列可插入到表达载体中形成重组表达载体中。术语“表达载体”指本领域熟知的细菌质粒、噬菌体、酵母质粒、植物细胞病毒、哺乳动物细胞病毒或其他载体。总之，只要能在宿主体内复制和稳定，任何质粒和载体都可以用。表达载体的一个重要特征是通常含有复制起点、启动子、标记基因和翻译控制元件。本发明中的表达载体不局限于下述实施例中提到的pYLCRISPR/Cas9载体。本领域的技术人员熟知的方法能用于构建含抗逆蛋白编码的核苷酸序列和合适的转录/翻译控制信号的表达载体。这些方法包括体外重组DNA技术、DNA合成技术、体内重组技术等。所述的DNA序列可有效连接到表达载体中的适当启动子上，以指导mRNA合成。表达载体还包括翻译起始用的核糖体结合位点和转录终止子。
本发明的含有a)所述多核苷酸的生物工程菌、或含有b)所述重组表达载体的生物工程菌，所述生物工程菌含有如上述所述的重组表达载体或基因组内整合有如第二方面所述的多核苷酸。生物工程菌以使其能够表达蛋白质。生物工程菌可以是原核细胞，如细菌细胞；或是低等真核细胞，如酵母细胞；或是高等真核细胞，如植物细胞。代表性例子有：大肠杆菌，链霉菌属、农杆菌；真菌细胞如酵母；植物细胞等。不局限于本申请中所采用的农杆菌Agl0。
本发明的含有a)所述多核苷酸的转基因植物细胞，或含有b)所述重组表达载体的转基因植物。本发明中，对于适用于本发明的植物没有特别的限制，只要其适合进行基因的转化操作既可以。
本发明第三方面，提供如第一方面所述蛋白作为靶标用于提高植物对重金属耐受性或提高植物重金属转运能力中的用途；优选包括促进植物生长，包括促进根系生长、提高花瓣的数量性状。本申请中，花瓣的数量性状包括花径、花型、花瓣大小和花瓣重量。
本发明第四方面，提供如第一方面所述蛋白或第二方面所述的生物材料筛选用于提高植物对重金属耐受性或提高植物重金属转运能力产品中的用途；优选包括促进植物生长，包括促进根系生长、提高花瓣的数量性状。
本发明，采用CRISPR/Cas9基因编辑技术将第一方面所述的重金属转运蛋白PhHMA5II-1失活导入矮牵牛，形成矮牵牛突变体，经铜胁迫实验验证，该矮牵牛突变体的花瓣中的铁含量明显降低，铜和锰含量也有不同程度的下降；同时矮牵牛突变体的生长得到了促进，特别是花瓣的数量性状得到了提高，尤其是花瓣的重量。说明该基因其能提高植物体内重金属转运能力或提高植物对重金属耐受性，特别是提高铁和铜的转运能力，由此显示了其在抗重金属植物培育的过程中起到了重要的作用。
本发明第五方面，提供一种用于提高植物对重金属耐受性或提高植物重金属转运能力的产品，包括PhHMA5II-1抑制剂或PhHMA5II-1基因干扰核酸构建体；
所述PhHMA5II-1抑制剂是指使第一方面所述的蛋白PhHMA5II-1质突变或失活或功能下降的化合物；
或，所述PhHMA5II-1基因干扰核酸构建体是指使第一方面所述的蛋白PhHMA5II-1质突变或失活或功能下降的编辑系统。
本发明的一个具体实施方式中，所述PhHMA5II-1基因干扰核酸构建体为：1)选择PhHMA5II-1基因的编码区如gtcaAGAAACTTTGCAGTTGCCA所示序列(SEQ ID NO.10)和如attgGCTACTCTGATTACAGAGG所示序列(SEQ ID NO.12)，作为CRISPR/Cas9系统的靶序列；2)将所述靶序列与载体片段pYLCRISPR/Cas9进行连接，得到所述PhHMA5II-1基因干扰核酸构建体。
本发明第六方面，提供一种构建转基因植物的方法，所述方法包括：向植物导入第五方面所述的PhHMA5II-1抑制剂或PhHMA5II-1基因干扰核酸构建体，得到植物突变体。
当采用重组DNA技术转化生物工程菌时，可用本领域技术人员熟知的常规技术进行。当生物工程菌为原核生物如大肠杆菌时，能吸收DNA的感受态细胞可在指数生长期后收获，用CaCl2法处理，所用的步骤在本领域众所周知。另一种方法是使用MgCl2。如果需要，转化也可用电穿孔的方法进行。当宿主是真核生物，可选用如下的DNA转染方法：磷酸钙共沉淀法，常规机械方法如显微注射、电穿孔、脂质体包装等。转化植物也可使用农杆菌转化或基因枪转化等方法，例如叶盘法、幼胚转化法等。对于转化的植物细胞、组织或器官可以用常规方法再生成植株，从而获得性状发生改变的植物。
本发明中，采用农杆菌介导法将如上述所述的多核苷酸或如上述所述的重组表达载体转入植物中。较佳的，所述农杆菌介导法为：将所述的重组表达载体转化于农杆菌；将含有所述重组表达载体的农杆菌转化于所述植物。
在本发明的一个具体实施例中，构建植物突变体的方法，包括1)选择PhHMA5II-1基因的编码区如gtcaAGAAACTTTGCAGTTGCCA所示序列(SEQ ID NO.10)和如attgGCTACTCTGATTACAGAGG所示序列(SEQ ID NO.12)，作为CRISPR/Cas9系统的靶序列；2)将所述靶序列与载体片段pYLCRISPR/Cas9进行连接，得到重组质粒；3)用含有所述重组质粒的农杆菌Agl 0浸染矮牵牛的愈伤组织，通过筛选，获得所述植物突变体。
本发明第七方面，提供一种培育对重金属耐受性高或对重金属转运能力高的植物的方法，在铜胁迫下，培育如第四方面得到的转基因植物。本发明中，所述铜为硫酸铜；所述硫酸铜的浓度不高于10μM。
本发明中，所述重金属包括铜、锌、锰和铁。较佳的，所述重金属为铜和铁。尤其是铁。
本发明中，对于适用于本发明的植物没有特别的限制，只要其适合进行基因的转化操作既可以。所述的植物不限于双子叶植物或单子叶植物。更具体地，所述双子叶植物包括拟南芥、紫花苜宿、矮牵牛、棉花、矮牵牛和蓖麻。在本发明的一个具体实施例中，所述植物优选矮牵牛。
本申请的下述实施例中，矮牵牛的品种为W115。
本申请的下述实施例中，LB固体培养基的配方为：10g胰蛋白胨(购于oxoid)，5g酵母粉(购于oxoid)，10g氯化钠(购于国药)，10g琼脂粉(购于上海生工)，补水至1000mL。
本申请的下述实施例中，LB液体培养基的配方为：10g胰蛋白胨(购于oxoid)，5g酵母粉(购于oxoid)，10g氯化钠(购于国药)，补水至1000mL。
本申请的下述实施例中，固体MS共培养基的配方为：5g蔗糖，2.5g葡萄糖，2g琼脂，1.1g MS，250μL 1mg/mL叶酸，250μL 2mg/mL 6-苄基腺嘌呤，250μL 0.1mg/mL 1-萘乙酸，250μL 1mg/mL玉米素，250μL 200mM乙酰丁香酮，加水至250mL，调节pH至5.8。
本申请的下述实施例中，MS固体选择培养基的配方为：5g蔗糖，2.5g葡萄糖，2g琼脂，1.1g MS，500μL 1mg/mL玉米素，250μL 1mg/mL叶酸，50mg/L卡那霉素，250mg/L羧苄青霉素，加水至250mL，调节pH至5.8。
本申请的下述实施例中，生根培养基的配方为：5g蔗糖，2.5g葡萄糖，2g琼脂,1.1gMS，250μL 1mg/mL叶酸，250mg/L羧苄青霉素，加水至250mL，调节pH至5.8。
本申请的下述实施例中，农杆菌菌株Agl0。
实施例1金属转运PhHMA5II-1基因的克隆
本实施例中，进行矮牵牛PhHMA5II-1基因的克隆，包括如下：
1.1植物材料的获得
取矮牵牛W115的叶片组织，用于提取RNA。
1.2总RNA的提取
取上述叶片组织，冻存后研磨得到粉末，然后与1.5mL的RNAiso Plus(Takara)在离心管混合，静置5min。
在上述离心管中加入300μL氯仿充分混匀，室温静置2分钟，于13200rpm离心15分钟，取800μL上清液转移至新的离心管中。加入800μL异丙醇(麦克林)，充分混匀，静置10分钟，于13200rpm离心10分钟，弃去上清。加1.5mL的70％酒精，涡旋10分钟，于13200rpm离心10分钟，弃去上清，空离2分钟，吸尽酒精。加入20μL～50μL的RNase FreeddH2O(生工生物工程(上海)股份有限公司)，室温30分钟，得到植物总RNA。
采用NanoDrop微量分光光度计，检测提取的RNA浓度。
1.3cDNA的获得
以1μg上述得到的总RNA为模板，进行cDNA的合成。采用PrimeScriptTM RT reagentKit with gDNA Eraser(Perfect Real Time)试剂盒进行反转录，操作参考试剂盒说明书。
引物：5'-GCGAGCACAGAATTAATACGACTCACTATAGGTTTTTTTTTTTTTTT-3'(SEQ IDNO.3)
1.4获得PhHMA5II-1基因的3'端基因序列
现有技术中仅对矮牵牛品种Petunia axillaris和Petunia inflata的基因组进行了测序，而对矮牵牛品种W115的基因组尚未测序。且矮牵牛品种Petunia axillaris和Petunia inflata基因组的3'端注释存在错误，为了得到矮牵牛W115中PhHMA5II-1基因的序列全长，通过3’RACE获得正确的矮牵牛PhHMA5II-1基因3'端基因序列。
以2.5μL的步骤1.3)得到的cDNA为模板，利用下述中的第一轮PCR引物，按照PrimeSTAR Max DNA Polymerase(Takara)试剂盒体系进行第一轮PCR扩增，反应条件如下：98℃预变性60s，接着32个循环：98℃变性10s；58℃退火20s；72℃延伸30s，之后72℃后延伸10min，获得第一轮PCR产物。
3'RACE PCR的第一轮PCR的引物序列为：
F:5'-CAAGGCAATTGTCGAATATGCC-3'(SEQ ID NO.4)
R:5'-GCGAGCACAGAATTAATACGACT-3'(SEQ ID NO.5)
将上述第一轮PCR产物用水稀释20倍，取1μL为模板，利用下述中的第二轮PCR引物，按照PrimeSTAR Max DNA Polymerase(Takara)试剂盒体系进行第二轮PCR扩增，反应条件为：98℃预变性60s，接着32个循环：98℃变性10s；55℃退火20s；72℃延伸20s，之后72℃后延伸10min，获得第二轮PCR产物。将获得的第二轮PCR产物送苏州金唯智公司进行测序。
3'RACE PCR的第二轮PCR的引物序列为：
F:5'-CTGTTTCAGCTGATGCCGGT-3'(SEQ ID NO.6)
R:5'-CGCGGATCCGAATTAATACGACTCACTATAGG-3'(SEQ ID NO.7)
1.5获得PhHMA5II-1全长基因
根据步骤1.4)中的测序结果，设计PhHMA5II-1全长基因的引物的序列如下：
正向引物
5'-GGGGACAAGTTTGTACAAAAAAGCAGGCTCAATGGCAACTGCAAAGTTTCTTTC-3'(SEQ IDNO.8)
反向引物
5'-GGGGACCACTTTGTACAAGAAAGCTGGGTTCACTCAACAGTTATGCCTCCAATC-3'(SEQ IDNO.9)
以上述步骤1.3)获得的cDNA为模板，用如SEQ ID NO.8所示引物和SEQ ID NO.9所示引物，按照PrimeSTAR Max DNA Polymerase(Takara)试剂盒体系进行PhHMA5II-1基因全长的扩增，得到长度为2973bp的PhHMA5II-1基因，序列如SEQ ID NO.1所示；根据开放读码框序列推导出共990个氨基酸的PhHMA5II-1蛋白，序列如SEQ ID NO.2所示。
上述PCR扩增的程序为：在PCR仪上，94℃预变性2分钟，94℃变性30秒，58℃退火30秒，72℃延伸1分钟，反应35个循环；最后72℃延伸7分钟，4℃暂存。
序列通过在NCBI网站进行BLASTP(http：//blast.ncbi.nlm.nih.gov/)比对已有的数据库(Genbank)，比较结果见图1。将PhHMA5II-1蛋白的氨基酸序列，采用参考文献1(Nuria Andrés-Colás,Vicente Sancenón,Susana Rodríguez-Navarro,et.TheArabidopsis heavy metal P-type ATPase HMA5 interacts with metallochaperonesand functions in copper detoxification of roots.The plant journal,45:225-236(2005))中记载，进行功能结构区域比较，比较结果见图2。
图1为本实施例扩增得到PhHMA5II-1的氨基酸序列和拟南芥AtHMA5氨基酸序列的同源比较结果图。
从图1可知，发现矮牵牛PhHMA5II-1的氨基酸序列与拟南芥AtHMA5氨基酸序列的序列一致性为73.5％。
图2为本实施例扩增得到PhHMA5II-1蛋白和拟南芥AtHMA5蛋白的氨基酸序列的功能结构域的比较结果图。
从图2可知，PhHMA5II-1蛋白含有HMA5家族蛋白含有的典型的保守结构域，如两个金属离子结合位点，1个ATP结合位点和8个跨膜结构域等。
实施例2 pYLCRISPR/Cas9的质粒构建和转化
本实施例中，将实施例1获得的PhHMA5II-1经基因编辑后与pYLCRISPR/Cas9连接，并转化于农杆菌中。包括如下：
2.1获得重组质粒
通过CRISPR-P 2.0和CRISPR-GE在线软件选取PhHMA5II-1基因敲除的两个靶序列，并通过CRISPR-GE在线软件设计相应的引物。参考文献2(Ma,X.et al.A RobustCRISPR/Cas9System for Convenient,High-Efficiency Multiplex Genome Editing inMonocot and Dicot Plants.Mol.Plant 8,1274–1284(2015))中记载的pYLCRISPR/Cas9进行酶切和连接的条件进行。
靶序列1由AtU3b启动子驱动表达，其用于构建载体的序列为：
正链：gtcaAGAAACTTTGCAGTTGCCA(SEQ ID NO.10)
负链：aaacTGGCAACTGCAAAGTTTCT(SEQ ID NO.11)
其中，小写字母gtca和aaac代表接头。
靶序列2由AtU6-1启动子驱动表达，其用于构建载体的序列为：
正链：attgGCTACTCTGATTACAGAGG(SEQ ID NO.12)
负链：aaacCCTCTGTAATCAGAGTAGC(SEQ ID NO.13)
其中，小写字母attg和aaac代表接头。
将靶序列1和靶序列2同时连入pYLCRISPR/Cas9载体，得到重组质粒，重组质粒的图谱如图3所示。
2.2重组质粒转化于农杆菌菌株中
取1μL步骤2.1得到的重组质粒与10μL农杆菌菌株Agl 0小心混匀，用电转仪(eppendorf)在1500v进行转化。
取30μL复苏液体涂布于含有150μg/mL卡那霉素(Kanamycine)的LB固体培养基上，并在28℃培养48h。
培养过夜后，挑取单菌落进行菌落PCR，甘油保存阳性单克隆菌落，放入-80℃冰箱保存。
实施例3将重组质粒转入模式植物矮牵牛
本实施例中，采用实施例2获得的阳性单克隆菌落侵染叶片，并培养，得到矮牵牛突变体，包括如下：
(1)预摇农杆菌
吸取40μL的阳性单克隆菌落至7mL的LB液体培养基中，并在28℃于200rpm摇菌培养12～16h；培养至吸光度OD600＝1.5～2.0，得到预摇农杆菌。预摇农杆菌中加入水至35mL并加3.5μL的200mM乙酰丁香酮，混匀，得到菌液。
(2)叶片灭菌
取矮牵牛“W115”的绿色无黄枯叶片，用70％酒精消毒15s，用0.4％次氯酸钠消毒10min，灭菌水泡2min，重复5次。
(3)转化植株：
配制35mL含有乙酰丁香酮的水溶液，其中水溶液中含有3.5μL的200mM乙酰丁香酮，用含有乙酰丁香酮的水溶液重悬预摇农杆菌。将灭菌后的叶片浸泡于前述重悬预摇农杆菌的溶液中15分钟。
(4)共培养：将浸泡后的叶片置于MS共培养基上，并在人工气候箱中黑暗培养48小时。
(5)愈伤组织和幼苗的诱导：共培养好的叶片移入含有MS固体选择培养基中；每隔2-3周将叶片转移至新的MS固体培养基中。
(6)生根：待愈伤转化成长茎的芽后，将茎切下，移入生根培养基中，待植株长出根后移入土中，培养，得到转化后的植株。
(7)突变体株系的鉴定
对上述转化后的植株提取DNA，采用如SEQ ID NO.14所示序列和SEQ ID NO.15所示序列的引物筛选获得阳性基因矮牵牛突变体。
引物F：ATGTTGACCGGTAAGGCGCG(SEQ ID NO.14)
引物R：AAACTTGCAGAATGGCTAGAGTCA(SEQ ID NO.15)
然后，以筛选得到的阳性植物的DNA为模板，采用如SEQ ID NO.16所示序列和SEQID NO.17所示序列的引物，按Master Mix(With Dye)(翌圣)试剂盒体系进行PCR，获得的PCR扩增产物用如SEQ ID NO.18所示序列的引物进行测序。PCR反应程序为：98℃预变性30s，接着32个循环：98℃变性10s；58℃退火30s；72℃延申30s，之后72℃后延伸10min。
引物F：CCATAGTCGGTTGATCAAATGT(SEQ ID NO.16)
引物R：ATGAGCACTGCTATTCAAGAATC(SEQ ID NO.17)
进行PCR，对PCR扩增产物，用引物CAACAACAGCTTCTTTAATACCAG(SEQ ID NO.18)进行测序，找出阳性矮牵牛突变体的突变位点。
图4为本实施例的矮牵牛突变体和野生型矮牵牛的部分基因序列对比图。图中-代表删除一个C碱基。
从图4可知，已成功获得矮牵牛突变体。
实施例4矮牵牛突变体和野生型矮牵牛的高铜胁迫考察
本实施例中，将实施例3获得的矮牵牛突变体进行自交，自交后的种子进行高铜胁迫考察，包括如下：
野生型矮牵牛W115种子(记为CK)以及矮牵牛突变体(记为PhHMA5II-1)的种子用0.08％次氯酸钠、70％酒精，涡旋仪振荡10min后，用灭菌水浸泡1min，重复5次，加入灭菌水后，4℃黑暗培养2d；然后将消毒后的种子分别播种在MS培养基上生长14d，生长期间给予光照16h，黑暗8h，温度25℃，得到幼苗。
用海绵条包裹幼苗茎部，每条海绵包裹一株。将包裹好的幼苗塞进泡沫浮板的圆孔中，在水培盒中用1/2Hoagland营养液培养28天。28天后，将幼苗移至由10μM硫酸铜和1/2Hoagland营养液组成的溶液中继续培养3～4周，期间收集新长出的花瓣。植物的生长条件为：在温度25℃和湿度为65％，光照和黑暗交替进行，光照时间为16h和黑暗时间为8h。用KOH将营养液的pH值调节至5.5，每7天更换一次。
每三朵花为一个重复样品，收集好的花瓣于105℃杀青30min，80℃过夜烘干至恒重，用于后续金属离子浓度测定。
称取烘干后的植物材料，与硝酸和30％过氧化氢(1：1)在120℃消解1.5h，用去离子水定容。用电感耦合等离子体发射光谱仪Optima 8000测定花瓣中的铜、锌、锰和铁的离子浓度。
图5为本实施例在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣中铜离子、锰离子和锌离子的含量图。
图6为本实施例在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣中铁离子的含量图。
图7为本实施例在10μM硫酸铜胁迫下，野生型矮牵牛和矮牵牛突变体的花瓣干重图。
从图5可知，铜胁迫后，与野生型矮牵牛相比，矮牵牛突变体的花瓣中铜和锰均有所下降，锌含量有所升高，但是均差异不显著。
从图6可知，矮牵牛突变体的花瓣中铁离子的含量，显著低于野生型矮牵牛。
从图7可知，矮牵牛突变体的花瓣的干重，显著高于野生型矮牵牛。
图5、图6和图7说明，本申请的重金属转运蛋白能提高植物体内重金属转运能力或提高植物对重金属耐受性，尤其是提高铁和铜的转运能力，特别是铁的转运能力，此外还能促进植物生长，提高花瓣数量性状，尤其是花瓣的干重。究其原因，可能是在高铜胁迫下，重金属转运蛋白能通过降低植物花瓣中的铁含量，来降低铜胁迫带来的负面影响，平衡重金属的浓度，从而促进植物生长，尤其是提高花瓣数量性状，特别是提高花瓣的干重。
以上的实施例是为了说明本发明公开的实施方案，并不能理解为对本发明的限制。此外，本文所列出的各种修改以及发明中方法、组合物的变化，在不脱离本发明的范围和精神的前提下对本领域内的技术人员来说是显而易见的。虽然已结合本发明的多种具体优选实施例对本发明进行了具体的描述，但应当理解，本发明不应仅限于这些具体实施例。事实上，各种如上所述的对本领域内的技术人员来说显而易见的修改来获取发明都应包括在本发明的范围内。
序列表
<110> 上海交通大学
<120> 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途
<160> 18
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2973
<212> DNA
<213> SEQ ID NO：1(Artificial Sequence)
<400> 1
atggcaactg caaagtttct ttcatttgca tgcataagaa atgaaagtaa atatggagat 60
ttttcatcaa gaccacatta tccatcaatg ccaaaatatc caaagggtgt tactatatca 120
actgatgaag aaaaaagtat acatggaaca gaatcaaagg caatatattc agtacatgga 180
atgagttgtt ctgcatgtgc aggttctgta gagaaagcaa ttaaaagact tgctggtatt 240
aaagaagctg ttgttgatgt cttgaataat agagctcaag ttatttttta tcctagctat 300
gttaatgagg aaatgatccg tgagaccata gaagatgttg gattccaggc tactctgatt 360
acagaggagg caacggagaa atcctcccaa gtctgccgga tacgtataaa aggaatgacc 420
tgcacttctt gctccacaac tgttgaatct gctttacagt tgatcccagg tgtacagaaa 480
gcacaagtag cattagcaac tgaagaggct gaaattcaat atgatccaag gatattaact 540
tacaatcagc ttttagagac catagaaaat actggatttg aagccatatt aattagcaca 600
ggagaagata ggagcaaaat attgctgaaa attgatggag tacacactgg cgattccatg 660
aggatcattg aaagttctct tagagcactc ccgggtgttg aagacattga tattgatcct 720
gagttgaaga agttatccgt ttcttataaa tcagatataa taggacctag agatttcatt 780
cgagtaattg agtcaactgg ttccggacag ttcaatgcaa tgttatatcc tgaaggaggt 840
ggggaacaat cgcatagaca ggaggaaatt ggaaactatc gtcggtcctt tctctggagc 900
ttgtttttta ccattccagt tttcttgact tccatggtct ttatgtatat tcctggtctg 960
aaggatggat tggatattaa agttgttaac atgcttagca ttggagagat tttgcggtgg 1020
gtgctttcaa ctccagtgca attcatcatt ggccgtcgtt tctattctgg ttcttacaaa 1080
gcattacgtc atggttctgc aaatatggat gttttgattg ctttgggaac aaatgcggcc 1140
tacttttatt cagtctactc ggtgttgaga gctgctactt ccccaagctt caagtctact 1200
gatttttttg agaccagctc aatgctcatt tccttcattc tactcgggaa gtatcttgaa 1260
gttttggcca aagggaagac atcggaggcc attgctaaac tcatgaactt gacccctgat 1320
actgcaacat tattacagca tggtgatgaa ggaaatgtgg taaatgagga agaaattgat 1380
agccgactga tacagaagaa tgatgtgatc agaatcgttc caggtgcaaa agtagcctgt 1440
gatggttttg tcatctgggg acaaagccac gtcaacgaga gtatgataac cggagaatct 1500
cggccagtgg ccaaacggaa gggcgacata gtgattggag gaactgtgaa tgagaatggt 1560
gttctgcata ttagagctac tagagttgga tcagagagtg cgctttcaca gattgttagg 1620
ctggttgaat cagcacagat ggctaaagct cctgttcaaa aatttgctga tggcatttct 1680
aaatattttg tgcctcttgt tattattgtc tccttttcca cttggctagc ctggttttta 1740
gctgggaaat acgccgccta tcctaaatcc tggattccat cttccatgga tagctttcag 1800
cttgccttgc aatttggtat atctgttatg gtcatagctt gcccgtgtgc ccttggtctt 1860
gctactccaa ctgctgtcat ggttggcaca ggagttggtg cttctcgtgg tgtcctgatt 1920
aaaggtgggc aggctttgga aagcgctcaa aaggtgaact gtatagtgtt tgataagacg 1980
ggtacactta caatggggaa accagtcgtt gtaaatacaa ggcttttcag aactatggta 2040
cttcgagaat tttatgaatt ggttgctgca gctgaggtaa atagtgagca ccccttagcc 2100
aaggcaattg tcgaatatgc caaaaagttt agagaggatg aggagaaccc ggtatggcct 2160
gaaatccagg actttgagtc aataactggc catggcgtga aggctattgt ccacagcaag 2220
aaagtaatag ttggaaacaa gagcttgatg ttagcacaag gcatttctgt ttcagctgat 2280
gccgatgaag ttctagctga aacagaagaa ttggctcaaa ctggaatact tgtatcagtt 2340
gatggtgtac tgactggagt tgttgctata tcagatcctg tgaagcccgg agctcgtgaa 2400
gtcatttctc ttcttaagtc tatgaatgtc gagagtaaac tagtaacagg tgacaattgg 2460
ggaacagcta atgccattgc caaggaagtt ggcattacgg atgttattgc agaagcaaaa 2520
cctgaagaca aagcagaaaa agtgaaggaa ttgcagagtt caggaaaagt tgttgcaatg 2580
gtcggagatg gaattaacga ttctccagct ctcgtagcag cagatgttgg aatggcaata 2640
ggggcgggga cagatatagc tattgaggca gctgatattg tcctaatgaa aagcaatctg 2700
gaagatgtca taactgctat tgacctttcc aggaaaacat ttggtcgaat tcgtctgaac 2760
tacttttggg catttggcta taaccttctt ggcatcccaa ttgctgctgg agctcttttc 2820
ccatctacta gatttcgcct gccaccgtgg gtagcaggtg cagcaatggc tgcttcttca 2880
gtgagtgttg tctgcagctc tcttttgttg aagaattaca cgagaccaaa gaagcttgat 2940
aatcttggga ttggaggcat aactgttgag tga 2973
<210> 2
<211> 990
<212> PRT
<213> SEQ ID NO：2(Artificial Sequence)
<400> 2
Met Ala Thr Ala Lys Phe Leu Ser Phe Ala Cys Ile Arg Asn Glu Ser
1 5 10 15
Lys Tyr Gly Asp Phe Ser Ser Arg Pro His Tyr Pro Ser Met Pro Lys
20 25 30
Tyr Pro Lys Gly Val Thr Ile Ser Thr Asp Glu Glu Lys Ser Ile His
35 40 45
Gly Thr Glu Ser Lys Ala Ile Tyr Ser Val His Gly Met Ser Cys Ser
50 55 60
Ala Cys Ala Gly Ser Val Glu Lys Ala Ile Lys Arg Leu Ala Gly Ile
65 70 75 80
Lys Glu Ala Val Val Asp Val Leu Asn Asn Arg Ala Gln Val Ile Phe
85 90 95
Tyr Pro Ser Tyr Val Asn Glu Glu Met Ile Arg Glu Thr Ile Glu Asp
100 105 110
Val Gly Phe Gln Ala Thr Leu Ile Thr Glu Glu Ala Thr Glu Lys Ser
115 120 125
Ser Gln Val Cys Arg Ile Arg Ile Lys Gly Met Thr Cys Thr Ser Cys
130 135 140
Ser Thr Thr Val Glu Ser Ala Leu Gln Leu Ile Pro Gly Val Gln Lys
145 150 155 160
Ala Gln Val Ala Leu Ala Thr Glu Glu Ala Glu Ile Gln Tyr Asp Pro
165 170 175
Arg Ile Leu Thr Tyr Asn Gln Leu Leu Glu Thr Ile Glu Asn Thr Gly
180 185 190
Phe Glu Ala Ile Leu Ile Ser Thr Gly Glu Asp Arg Ser Lys Ile Leu
195 200 205
Leu Lys Ile Asp Gly Val His Thr Gly Asp Ser Met Arg Ile Ile Glu
210 215 220
Ser Ser Leu Arg Ala Leu Pro Gly Val Glu Asp Ile Asp Ile Asp Pro
225 230 235 240
Glu Leu Lys Lys Leu Ser Val Ser Tyr Lys Ser Asp Ile Ile Gly Pro
245 250 255
Arg Asp Phe Ile Arg Val Ile Glu Ser Thr Gly Ser Gly Gln Phe Asn
260 265 270
Ala Met Leu Tyr Pro Glu Gly Gly Gly Glu Gln Ser His Arg Gln Glu
275 280 285
Glu Ile Gly Asn Tyr Arg Arg Ser Phe Leu Trp Ser Leu Phe Phe Thr
290 295 300
Ile Pro Val Phe Leu Thr Ser Met Val Phe Met Tyr Ile Pro Gly Leu
305 310 315 320
Lys Asp Gly Leu Asp Ile Lys Val Val Asn Met Leu Ser Ile Gly Glu
325 330 335
Ile Leu Arg Trp Val Leu Ser Thr Pro Val Gln Phe Ile Ile Gly Arg
340 345 350
Arg Phe Tyr Ser Gly Ser Tyr Lys Ala Leu Arg His Gly Ser Ala Asn
355 360 365
Met Asp Val Leu Ile Ala Leu Gly Thr Asn Ala Ala Tyr Phe Tyr Ser
370 375 380
Val Tyr Ser Val Leu Arg Ala Ala Thr Ser Pro Ser Phe Lys Ser Thr
385 390 395 400
Asp Phe Phe Glu Thr Ser Ser Met Leu Ile Ser Phe Ile Leu Leu Gly
405 410 415
Lys Tyr Leu Glu Val Leu Ala Lys Gly Lys Thr Ser Glu Ala Ile Ala
420 425 430
Lys Leu Met Asn Leu Thr Pro Asp Thr Ala Thr Leu Leu Gln His Gly
435 440 445
Asp Glu Gly Asn Val Val Asn Glu Glu Glu Ile Asp Ser Arg Leu Ile
450 455 460
Gln Lys Asn Asp Val Ile Arg Ile Val Pro Gly Ala Lys Val Ala Cys
465 470 475 480
Asp Gly Phe Val Ile Trp Gly Gln Ser His Val Asn Glu Ser Met Ile
485 490 495
Thr Gly Glu Ser Arg Pro Val Ala Lys Arg Lys Gly Asp Ile Val Ile
500 505 510
Gly Gly Thr Val Asn Glu Asn Gly Val Leu His Ile Arg Ala Thr Arg
515 520 525
Val Gly Ser Glu Ser Ala Leu Ser Gln Ile Val Arg Leu Val Glu Ser
530 535 540
Ala Gln Met Ala Lys Ala Pro Val Gln Lys Phe Ala Asp Gly Ile Ser
545 550 555 560
Lys Tyr Phe Val Pro Leu Val Ile Ile Val Ser Phe Ser Thr Trp Leu
565 570 575
Ala Trp Phe Leu Ala Gly Lys Tyr Ala Ala Tyr Pro Lys Ser Trp Ile
580 585 590
Pro Ser Ser Met Asp Ser Phe Gln Leu Ala Leu Gln Phe Gly Ile Ser
595 600 605
Val Met Val Ile Ala Cys Pro Cys Ala Leu Gly Leu Ala Thr Pro Thr
610 615 620
Ala Val Met Val Gly Thr Gly Val Gly Ala Ser Arg Gly Val Leu Ile
625 630 635 640
Lys Gly Gly Gln Ala Leu Glu Ser Ala Gln Lys Val Asn Cys Ile Val
645 650 655
Phe Asp Lys Thr Gly Thr Leu Thr Met Gly Lys Pro Val Val Val Asn
660 665 670
Thr Arg Leu Phe Arg Thr Met Val Leu Arg Glu Phe Tyr Glu Leu Val
675 680 685
Ala Ala Ala Glu Val Asn Ser Glu His Pro Leu Ala Lys Ala Ile Val
690 695 700
Glu Tyr Ala Lys Lys Phe Arg Glu Asp Glu Glu Asn Pro Val Trp Pro
705 710 715 720
Glu Ile Gln Asp Phe Glu Ser Ile Thr Gly His Gly Val Lys Ala Ile
725 730 735
Val His Ser Lys Lys Val Ile Val Gly Asn Lys Ser Leu Met Leu Ala
740 745 750
Gln Gly Ile Ser Val Ser Ala Asp Ala Asp Glu Val Leu Ala Glu Thr
755 760 765
Glu Glu Leu Ala Gln Thr Gly Ile Leu Val Ser Val Asp Gly Val Leu
770 775 780
Thr Gly Val Val Ala Ile Ser Asp Pro Val Lys Pro Gly Ala Arg Glu
785 790 795 800
Val Ile Ser Leu Leu Lys Ser Met Asn Val Glu Ser Lys Leu Val Thr
805 810 815
Gly Asp Asn Trp Gly Thr Ala Asn Ala Ile Ala Lys Glu Val Gly Ile
820 825 830
Thr Asp Val Ile Ala Glu Ala Lys Pro Glu Asp Lys Ala Glu Lys Val
835 840 845
Lys Glu Leu Gln Ser Ser Gly Lys Val Val Ala Met Val Gly Asp Gly
850 855 860
Ile Asn Asp Ser Pro Ala Leu Val Ala Ala Asp Val Gly Met Ala Ile
865 870 875 880
Gly Ala Gly Thr Asp Ile Ala Ile Glu Ala Ala Asp Ile Val Leu Met
885 890 895
Lys Ser Asn Leu Glu Asp Val Ile Thr Ala Ile Asp Leu Ser Arg Lys
900 905 910
Thr Phe Gly Arg Ile Arg Leu Asn Tyr Phe Trp Ala Phe Gly Tyr Asn
915 920 925
Leu Leu Gly Ile Pro Ile Ala Ala Gly Ala Leu Phe Pro Ser Thr Arg
930 935 940
Phe Arg Leu Pro Pro Trp Val Ala Gly Ala Ala Met Ala Ala Ser Ser
945 950 955 960
Val Ser Val Val Cys Ser Ser Leu Leu Leu Lys Asn Tyr Thr Arg Pro
965 970 975
Lys Lys Leu Asp Asn Leu Gly Ile Gly Gly Ile Thr Val Glu
980 985 990
<210> 3
<211> 47
<212> DNA
<213> SEQ ID NO：3(Artificial Sequence)
<400> 3
gcgagcacag aattaatacg actcactata ggtttttttt ttttttt 47
<210> 4
<211> 22
<212> DNA
<213> SEQ ID NO：4(Artificial Sequence)
<400> 4
caaggcaatt gtcgaatatg cc 22
<210> 5
<211> 23
<212> DNA
<213> SEQ ID NO：5(Artificial Sequence)
<400> 5
gcgagcacag aattaatacg act 23
<210> 6
<211> 20
<212> DNA
<213> SEQ ID NO：6(Artificial Sequence)
<400> 6
ctgtttcagc tgatgccggt 20
<210> 7
<211> 32
<212> DNA
<213> SEQ ID NO：7(Artificial Sequence)
<400> 7
cgcggatccg aattaatacg actcactata gg 32
<210> 8
<211> 54
<212> DNA
<213> SEQ ID NO：8(Artificial Sequence)
<400> 8
ggggacaagt ttgtacaaaa aagcaggctc aatggcaact gcaaagtttc tttc 54
<210> 9
<211> 54
<212> DNA
<213> SEQ ID NO.9(Artificial Sequence)
<400> 9
ggggaccact ttgtacaaga aagctgggtt cactcaacag ttatgcctcc aatc 54
<210> 10
<211> 23
<212> DNA
<213> SEQ ID NO.10(Artificial Sequence)
<400> 10
gtcaagaaac tttgcagttg cca 23
<210> 11
<211> 23
<212> DNA
<213> SEQ ID NO.11(Artificial Sequence)
<400> 11
aaactggcaa ctgcaaagtt tct 23
<210> 12
<211> 23
<212> DNA
<213> SEQ ID NO.12(Artificial Sequence)
<400> 12
attggctact ctgattacag agg 23
<210> 13
<211> 23
<212> DNA
<213> SEQ ID NO.13(Artificial Sequence)
<400> 13
aaaccctctg taatcagagt agc 23
<210> 14
<211> 20
<212> DNA
<213> SEQ ID NO.14(Artificial Sequence)
<400> 14
atgttgaccg gtaaggcgcg 20
<210> 15
<211> 24
<212> DNA
<213> SEQ ID NO.15(Artificial Sequence)
<400> 15
aaacttgcag aatggctaga gtca 24
<210> 16
<211> 22
<212> DNA
<213> SEQ ID NO.16(Artificial Sequence)
<400> 16
ccatagtcgg ttgatcaaat gt 22
<210> 17
<211> 23
<212> DNA
<213> SEQ ID NO.17(Artificial Sequence)
<400> 17
atgagcactg ctattcaaga atc 23
<210> 18
<211> 24
<212> DNA
<213> SEQ ID NO.18(Artificial Sequence)
<400> 18
caacaacagc ttctttaata ccag 24
Claims (11)
1.一种重金属转运蛋白PhHMA5II-1，其特征在于，所述蛋白为由如SEQ ID NO.2所示的氨基酸序列组成的蛋白。
2.与如权利要求1所述蛋白相关的生物材料，其特征在于，所述生物材料为如下任一种：
a)编码如权利要求1所述蛋白的多核苷酸；
b)含有a)所述多核苷酸的重组表达载体；
c)含有a)所述多核苷酸的生物工程菌，或含有b)所述重组表达载体的生物工程菌。
3.如权利要求2所述的生物材料，其特征在于，a)中，所述多核苷酸的核苷酸序列包括如SEQ ID NO.1所示序列。
4.如权利要求1所述的蛋白作为靶标用于提高植物对重金属耐受性或提高植物重金属转运能力中的用途，所述提高植物对重金属耐受性或提高植物重金属转运能力是通过使植物中所述蛋白失活或功能下降获得，所述植物为矮牵牛。
5.如权利要求4所述的用途，其特征在于，所述用途还包括促进植物生长和/或提高花瓣重量。
6.如权利要求1所述的蛋白或如权利要求2或3所述的生物材料在筛选用于提高植物对重金属耐受性或提高植物重金属转运能力产品中的用途，所述提高对重金属耐受性或提高植物重金属转运能力是通过使植物中所述蛋白失活或功能下降获得，所述植物为矮牵牛。
7.如权利要求6所述的用途，其特征在于，所述用途还包括促进植物生长和/或提高花瓣的重量。
8.一种用于提高植物对重金属耐受性或提高植物重金属转运能力的产品，其特征在于，所述产品为PhHMA5II-1基因干扰核酸构建体，所述PhHMA5II-1基因干扰核酸构建体是指使权利要求1所述的蛋白PhHMA5II-1突变或失活或功能下降的编辑系统，所述PhHMA5II-1基因干扰核酸构建体为：1)选择PhHMA5II-1基因的编码区如SEQ ID NO.10所示序列和如SEQ ID NO.12所示序列，作为CRISPR/Cas9系统的靶序列；2)将所述靶序列与载体片段pYLCRISPR/Cas9进行连接，得到所述PhHMA5II-1基因干扰核酸构建体。
9.一种制备植物突变体的方法，其特征在于，向植物导入权利要求8所述的产品，得到植物突变体，所述植物为矮牵牛。
10.一种培育对重金属耐受性高或对重金属转运能力高的植物的方法，其特征在于，在铜胁迫下，种植如权利要求9所述的方法得到的植物突变体，进行培育。
11.如权利要求4-7任一项所述的用途，其特征在于，所述重金属包括铜、锌、锰和铁。
Priority Applications (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN202111154498.3A CN114106120B (zh) | 2021-09-29 | 2021-09-29 | 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 |
Applications Claiming Priority (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN202111154498.3A CN114106120B (zh) | 2021-09-29 | 2021-09-29 | 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 |
Publications (2)
| Publication Number | Publication Date |
|---|---|
| CN114106120A CN114106120A (zh) | 2022-03-01 |
| CN114106120B true CN114106120B (zh) | 2023-09-08 |
Family
ID=80441497
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| CN202111154498.3A Active CN114106120B (zh) | 2021-09-29 | 2021-09-29 | 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 |
Country Status (1)
| Country | Link |
|---|---|
| CN (1) | CN114106120B (zh) |
Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO2012060705A1 (en) * | 2010-11-03 | 2012-05-10 | Vereniging Voor Christelijk Hoger Onderwijs, Wetenschappelijk Onderzoek En Patiënten Zorg | Plants with increased tolerance to metal ions |
| CN106349352A (zh) * | 2016-10-27 | 2017-01-25 | 上海交通大学 | 青蒿转运蛋白AaPDR3及其应用 |
Family Cites Families (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| KR100845279B1 (ko) * | 2006-11-28 | 2008-07-09 | 포항공과대학교 산학협력단 | 중금속이나 염 축적성, 또는 중금속, 염 또는 건조에 대한내성을 변화시키는 유전자 및 이들을 이용하여 제조한형질전환체 |
-
2021
- 2021-09-29 CN CN202111154498.3A patent/CN114106120B/zh active Active
Patent Citations (2)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO2012060705A1 (en) * | 2010-11-03 | 2012-05-10 | Vereniging Voor Christelijk Hoger Onderwijs, Wetenschappelijk Onderzoek En Patiënten Zorg | Plants with increased tolerance to metal ions |
| CN106349352A (zh) * | 2016-10-27 | 2017-01-25 | 上海交通大学 | 青蒿转运蛋白AaPDR3及其应用 |
Non-Patent Citations (1)
| Title |
|---|
| 蒺藜苜蓿重金属转运蛋白HMA5基因在共生结瘤中的功能分析;杨墨等;《科学通报》;20210511;第66卷(第Z2期);第3719-3731页 * |
Also Published As
| Publication number | Publication date |
|---|---|
| CN114106120A (zh) | 2022-03-01 |
Similar Documents
| Publication | Publication Date | Title |
|---|---|---|
| CN112626080B (zh) | 一种控制大豆-根瘤菌匹配性的r基因及其蛋白质和应用 | |
| CN111574605B (zh) | 水稻基因OsLAT5在调节敌草快的吸收积累中的应用 | |
| CN116179589B (zh) | SlPRMT5基因及其蛋白在调控番茄果实产量中的应用 | |
| CN107779456B (zh) | 蒺藜苜蓿MtWOX11基因及其在提高种子脂肪酸含量中的应用 | |
| CN110804090B (zh) | 蛋白质CkWRKY33及其编码基因与应用 | |
| CN113234751B (zh) | 基于根瘤菌III型效应因子NopP的农杆菌转化载体及其应用 | |
| CN104725496A (zh) | 一种调控植物开花的棉花WRKY转录因子GarWRKY9及应用 | |
| CN112048515B (zh) | 一种油菜S-腺苷-L-蛋氨酸依赖的甲基转移酶基因BnPMT6及其应用 | |
| CN109929019A (zh) | 一种与植物耐盐碱相关蛋白GsERF7及其编码基因与应用 | |
| CN110819634B (zh) | 一种细叶百合基因LpNAC6的克隆及其应用 | |
| CN109337884B9 (zh) | 一种丙酮酸激酶基因及其应用 | |
| CN108707614B (zh) | 一种花生抗逆性基因及其应用 | |
| CN114106120B (zh) | 一种重金属转运蛋白PhHMA5II-1及其相关产品和用途 | |
| CN113481176B (zh) | GA3ox1蛋白在调控苜蓿株型中的应用 | |
| CN115927237A (zh) | 一种油菜海藻糖-6-磷酸合成酶基因在调控含油量与脂肪酸组成中的用途 | |
| CN114213515B (zh) | 基因OsR498G0917707800.01及其编码的蛋白在调控水稻垩白中的应用 | |
| CN111560055B (zh) | 水稻基因OsLAT3在调节敌草快的吸收累积中的应用 | |
| CN112143737B (zh) | OsbZIP62-VP64融合表达改良水稻农艺性状的应用 | |
| CN110760522B (zh) | Ak209基因及编码蛋白与抗逆增产的应用 | |
| CN108728447B (zh) | 一种花生抗逆相关基因及其应用 | |
| CN110684749A (zh) | 玉米3-磷酸甘油脱氢酶ZmGPDH4及其编码基因在调控植物耐逆性中的应用 | |
| CN114717245B (zh) | MsbHLH35基因及其编码蛋白在调控紫花苜蓿产量和耐渍性中的应用 | |
| CN115011631B (zh) | 调控玉米苗期抗旱性的蛋白及其编码基因和应用 | |
| CN114958866B (zh) | 调控大豆分枝数的基因及其用途 | |
| CN113789327B (zh) | 一种铁皮石斛组成型强启动子ProDoWOX1及其应用 |
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 |