CN104098664B - 拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 - Google Patents
拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 Download PDFInfo
- Publication number
- CN104098664B CN104098664B CN201410040966.8A CN201410040966A CN104098664B CN 104098664 B CN104098664 B CN 104098664B CN 201410040966 A CN201410040966 A CN 201410040966A CN 104098664 B CN104098664 B CN 104098664B
- Authority
- CN
- China
- Prior art keywords
- ser
- glu
- lys
- leu
- val
- 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.)
- Expired - Fee Related
Links
- 108090000623 proteins and genes Proteins 0.000 title claims abstract description 73
- 230000017260 vegetative to reproductive phase transition of meristem Effects 0.000 title claims abstract description 26
- 230000033228 biological regulation Effects 0.000 title claims abstract description 16
- 102000000584 Calmodulin Human genes 0.000 title claims abstract description 15
- 108010041952 Calmodulin Proteins 0.000 title claims abstract description 15
- 241000219194 Arabidopsis Species 0.000 claims description 46
- 108091000084 calmodulin binding Proteins 0.000 claims description 20
- 230000014509 gene expression Effects 0.000 claims description 12
- KOSRFJWDECSPRO-UHFFFAOYSA-N alpha-L-glutamyl-L-glutamic acid Natural products OC(=O)CCC(N)C(=O)NC(CCC(O)=O)C(O)=O KOSRFJWDECSPRO-UHFFFAOYSA-N 0.000 claims description 9
- ZKAOJVJQGVUIIU-GUBZILKMSA-N Asp-Pro-Arg Chemical compound OC(=O)C[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCCNC(N)=N)C(O)=O ZKAOJVJQGVUIIU-GUBZILKMSA-N 0.000 claims description 6
- 241000880493 Leptailurus serval Species 0.000 claims description 6
- 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 claims description 6
- MQUZANJDFOQOBX-SRVKXCTJSA-N Ser-Phe-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CO)C(O)=O MQUZANJDFOQOBX-SRVKXCTJSA-N 0.000 claims description 6
- 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 claims description 6
- NLNCNKIVJPEFBC-DLOVCJGASA-N Val-Val-Glu Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@H](C(O)=O)CCC(O)=O NLNCNKIVJPEFBC-DLOVCJGASA-N 0.000 claims description 6
- 108010018691 arginyl-threonyl-arginine Proteins 0.000 claims description 6
- 108010062796 arginyllysine Proteins 0.000 claims description 6
- FSXRLASFHBWESK-UHFFFAOYSA-N dipeptide phenylalanyl-tyrosine Natural products C=1C=C(O)C=CC=1CC(C(O)=O)NC(=O)C(N)CC1=CC=CC=C1 FSXRLASFHBWESK-UHFFFAOYSA-N 0.000 claims description 6
- 108010055341 glutamyl-glutamic acid Proteins 0.000 claims description 6
- 108010050848 glycylleucine Proteins 0.000 claims description 6
- 108010034529 leucyl-lysine Proteins 0.000 claims description 6
- 108010017391 lysylvaline Proteins 0.000 claims description 6
- 108010056582 methionylglutamic acid Proteins 0.000 claims description 6
- 108010051242 phenylalanylserine Proteins 0.000 claims description 6
- 108010073969 valyllysine Proteins 0.000 claims description 6
- AWAXZRDKUHOPBO-GUBZILKMSA-N Ala-Gln-Lys Chemical compound C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCCN)C(O)=O AWAXZRDKUHOPBO-GUBZILKMSA-N 0.000 claims description 3
- BGNLUHXLSAQYRQ-FXQIFTODSA-N Ala-Glu-Gln Chemical compound C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O BGNLUHXLSAQYRQ-FXQIFTODSA-N 0.000 claims description 3
- HXNNRBHASOSVPG-GUBZILKMSA-N Ala-Glu-Leu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O HXNNRBHASOSVPG-GUBZILKMSA-N 0.000 claims description 3
- WGDNWOMKBUXFHR-BQBZGAKWSA-N Ala-Gly-Arg Chemical compound C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCN=C(N)N WGDNWOMKBUXFHR-BQBZGAKWSA-N 0.000 claims description 3
- MQIGTEQXYCRLGK-BQBZGAKWSA-N Ala-Gly-Pro Chemical compound C[C@H](N)C(=O)NCC(=O)N1CCC[C@H]1C(O)=O MQIGTEQXYCRLGK-BQBZGAKWSA-N 0.000 claims description 3
- RZZMZYZXNJRPOJ-BJDJZHNGSA-N Ala-Ile-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](C)N RZZMZYZXNJRPOJ-BJDJZHNGSA-N 0.000 claims description 3
- 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 claims 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 claims description 3
- CCDFBRZVTDDJNM-GUBZILKMSA-N Ala-Leu-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O CCDFBRZVTDDJNM-GUBZILKMSA-N 0.000 claims description 3
- MDNAVFBZPROEHO-DCAQKATOSA-N Ala-Lys-Val Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C(C)C)C(O)=O MDNAVFBZPROEHO-DCAQKATOSA-N 0.000 claims description 3
- MDNAVFBZPROEHO-UHFFFAOYSA-N Ala-Lys-Val Natural products CC(C)C(C(O)=O)NC(=O)C(NC(=O)C(C)N)CCCCN MDNAVFBZPROEHO-UHFFFAOYSA-N 0.000 claims description 3
- BGGAIXWIZCIFSG-XDTLVQLUSA-N Ala-Tyr-Glu Chemical compound [H]N[C@@H](C)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(O)=O)C(O)=O BGGAIXWIZCIFSG-XDTLVQLUSA-N 0.000 claims description 3
- SGYSTDWPNPKJPP-GUBZILKMSA-N Arg-Ala-Arg Chemical compound NC(=N)NCCC[C@H](N)C(=O)N[C@@H](C)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O SGYSTDWPNPKJPP-GUBZILKMSA-N 0.000 claims description 3
- FRMQITGHXMUNDF-GMOBBJLQSA-N Arg-Ile-Asn Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N FRMQITGHXMUNDF-GMOBBJLQSA-N 0.000 claims description 3
- UAOSDDXCTBIPCA-QXEWZRGKSA-N Arg-Ile-Gly Chemical compound CC[C@H](C)[C@@H](C(=O)NCC(=O)O)NC(=O)[C@H](CCCN=C(N)N)N UAOSDDXCTBIPCA-QXEWZRGKSA-N 0.000 claims description 3
- NIUDXSFNLBIWOB-DCAQKATOSA-N Arg-Leu-Cys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CS)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N NIUDXSFNLBIWOB-DCAQKATOSA-N 0.000 claims description 3
- UZGFHWIJWPUPOH-IHRRRGAJSA-N Arg-Leu-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N UZGFHWIJWPUPOH-IHRRRGAJSA-N 0.000 claims description 3
- KXOPYFNQLVUOAQ-FXQIFTODSA-N Arg-Ser-Ala Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CO)C(=O)N[C@@H](C)C(O)=O KXOPYFNQLVUOAQ-FXQIFTODSA-N 0.000 claims description 3
- OQPAZKMGCWPERI-GUBZILKMSA-N Arg-Ser-Val Chemical compound [H]N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(O)=O OQPAZKMGCWPERI-GUBZILKMSA-N 0.000 claims description 3
- XMGVWQWEWWULNS-BPUTZDHNSA-N Arg-Trp-Ser Chemical compound C1=CC=C2C(=C1)C(=CN2)C[C@@H](C(=O)N[C@@H](CO)C(=O)O)NC(=O)[C@H](CCCN=C(N)N)N XMGVWQWEWWULNS-BPUTZDHNSA-N 0.000 claims description 3
- POOCJCRBHHMAOS-FXQIFTODSA-N Asn-Arg-Ser Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CO)C(O)=O POOCJCRBHHMAOS-FXQIFTODSA-N 0.000 claims description 3
- IYVSIZAXNLOKFQ-BYULHYEWSA-N Asn-Asp-Val Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](C(C)C)C(O)=O IYVSIZAXNLOKFQ-BYULHYEWSA-N 0.000 claims description 3
- UBKOVSLDWIHYSY-ACZMJKKPSA-N Asn-Glu-Ser Chemical compound [H]N[C@@H](CC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O UBKOVSLDWIHYSY-ACZMJKKPSA-N 0.000 claims description 3
- OLVIPTLKNSAYRJ-YUMQZZPRSA-N Asn-Gly-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CC(=O)N)N OLVIPTLKNSAYRJ-YUMQZZPRSA-N 0.000 claims description 3
- JQSWHKKUZMTOIH-QWRGUYRKSA-N Asn-Gly-Phe Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CC(=O)N)N JQSWHKKUZMTOIH-QWRGUYRKSA-N 0.000 claims description 3
- 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 claims description 3
- RTFWCVDISAMGEQ-SRVKXCTJSA-N Asn-Phe-Asn Chemical compound C1=CC=C(C=C1)C[C@@H](C(=O)N[C@@H](CC(=O)N)C(=O)O)NC(=O)[C@H](CC(=O)N)N RTFWCVDISAMGEQ-SRVKXCTJSA-N 0.000 claims description 3
- AWXDRZJQCVHCIT-DCAQKATOSA-N Asn-Pro-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)CC(N)=O AWXDRZJQCVHCIT-DCAQKATOSA-N 0.000 claims description 3
- VPSHHQXIWLGVDD-ZLUOBGJFSA-N Asp-Asp-Asp Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O VPSHHQXIWLGVDD-ZLUOBGJFSA-N 0.000 claims description 3
- PXLNPFOJZQMXAT-BYULHYEWSA-N Asp-Asp-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](CC(O)=O)NC(=O)[C@@H](N)CC(O)=O PXLNPFOJZQMXAT-BYULHYEWSA-N 0.000 claims description 3
- AMRANMVXQWXNAH-ZLUOBGJFSA-N Asp-Cys-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](CS)NC(=O)[C@@H](N)CC(O)=O AMRANMVXQWXNAH-ZLUOBGJFSA-N 0.000 claims description 3
- 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 claims description 3
- VILLWIDTHYPSLC-PEFMBERDSA-N Asp-Glu-Ile Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O VILLWIDTHYPSLC-PEFMBERDSA-N 0.000 claims description 3
- OVPHVTCDVYYTHN-AVGNSLFASA-N Asp-Glu-Phe Chemical compound OC(=O)C[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 OVPHVTCDVYYTHN-AVGNSLFASA-N 0.000 claims description 3
- PZXPWHFYZXTFBI-YUMQZZPRSA-N Asp-Gly-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)CNC(=O)[C@@H](N)CC(O)=O PZXPWHFYZXTFBI-YUMQZZPRSA-N 0.000 claims description 3
- SVABRQFIHCSNCI-FOHZUACHSA-N Asp-Gly-Thr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)NCC(=O)N[C@@H]([C@@H](C)O)C(O)=O SVABRQFIHCSNCI-FOHZUACHSA-N 0.000 claims description 3
- SEMWSADZTMJELF-BYULHYEWSA-N Asp-Ile-Gly Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)NCC(O)=O SEMWSADZTMJELF-BYULHYEWSA-N 0.000 claims description 3
- SPWXXPFDTMYTRI-IUKAMOBKSA-N Asp-Ile-Thr Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O SPWXXPFDTMYTRI-IUKAMOBKSA-N 0.000 claims description 3
- DWOGMPWRQQWPPF-GUBZILKMSA-N Asp-Leu-Glu Chemical compound [H]N[C@@H](CC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O DWOGMPWRQQWPPF-GUBZILKMSA-N 0.000 claims description 3
- USENATHVGFXRNO-SRVKXCTJSA-N Asp-Tyr-Asp Chemical compound OC(=O)C[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](CC(O)=O)C(O)=O)CC1=CC=C(O)C=C1 USENATHVGFXRNO-SRVKXCTJSA-N 0.000 claims description 3
- VHUKCUHLFMRHOD-MELADBBJSA-N Asp-Tyr-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CC2=CC=C(C=C2)O)NC(=O)[C@H](CC(=O)O)N)C(=O)O VHUKCUHLFMRHOD-MELADBBJSA-N 0.000 claims description 3
- GGBQDSHTXKQSLP-NHCYSSNCSA-N Asp-Val-Lys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CC(=O)O)N GGBQDSHTXKQSLP-NHCYSSNCSA-N 0.000 claims description 3
- CWHKESLHINPNBX-XIRDDKMYSA-N Cys-Lys-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@@H](NC(=O)[C@@H](N)CS)CCCCN)C(O)=O)=CNC2=C1 CWHKESLHINPNBX-XIRDDKMYSA-N 0.000 claims description 3
- CNAMJJOZGXPDHW-IHRRRGAJSA-N Cys-Pro-Phe Chemical compound N[C@@H](CS)C(=O)N1CCC[C@H]1C(=O)N[C@@H](Cc1ccccc1)C(O)=O CNAMJJOZGXPDHW-IHRRRGAJSA-N 0.000 claims description 3
- MHYHLWUGWUBUHF-GUBZILKMSA-N Cys-Val-Arg Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)O)NC(=O)[C@H](CS)N MHYHLWUGWUBUHF-GUBZILKMSA-N 0.000 claims description 3
- KZEUVLLVULIPNX-GUBZILKMSA-N Gln-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)N)N KZEUVLLVULIPNX-GUBZILKMSA-N 0.000 claims description 3
- VVWWRZZMPSPVQU-KBIXCLLPSA-N Gln-Cys-Ile Chemical compound CC[C@H](C)[C@@H](C(=O)O)NC(=O)[C@H](CS)NC(=O)[C@H](CCC(=O)N)N VVWWRZZMPSPVQU-KBIXCLLPSA-N 0.000 claims description 3
- ZNZPKVQURDQFFS-FXQIFTODSA-N Gln-Glu-Ser Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O ZNZPKVQURDQFFS-FXQIFTODSA-N 0.000 claims description 3
- YPMDZWPZFOZYFG-GUBZILKMSA-N Gln-Leu-Ser Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CO)C(O)=O YPMDZWPZFOZYFG-GUBZILKMSA-N 0.000 claims description 3
- HHRAEXBUNGTOGZ-IHRRRGAJSA-N Gln-Phe-Gln Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O HHRAEXBUNGTOGZ-IHRRRGAJSA-N 0.000 claims description 3
- QENSHQJGWGRPQS-QEJZJMRPSA-N Gln-Ser-Trp Chemical compound C1=CC=C2C(C[C@H](NC(=O)[C@H](CO)NC(=O)[C@H](CCC(N)=O)N)C(O)=O)=CNC2=C1 QENSHQJGWGRPQS-QEJZJMRPSA-N 0.000 claims description 3
- OTQSTOXRUBVWAP-NRPADANISA-N Gln-Ser-Val Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(O)=O OTQSTOXRUBVWAP-NRPADANISA-N 0.000 claims description 3
- ICRKQMRFXYDYMK-LAEOZQHASA-N Gln-Val-Asn Chemical compound [H]N[C@@H](CCC(N)=O)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(N)=O)C(O)=O ICRKQMRFXYDYMK-LAEOZQHASA-N 0.000 claims description 3
- RUFHOVYUYSNDNY-ACZMJKKPSA-N Glu-Ala-Ala Chemical compound OC(=O)[C@H](C)NC(=O)[C@H](C)NC(=O)[C@@H](N)CCC(O)=O RUFHOVYUYSNDNY-ACZMJKKPSA-N 0.000 claims description 3
- 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 claims description 3
- OJGLIOXAKGFFDW-SRVKXCTJSA-N Glu-Arg-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)[C@H](CCCN=C(N)N)NC(=O)[C@H](CCC(=O)O)N OJGLIOXAKGFFDW-SRVKXCTJSA-N 0.000 claims description 3
- KEBACWCLVOXFNC-DCAQKATOSA-N Glu-Arg-Met Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCSC)C(O)=O KEBACWCLVOXFNC-DCAQKATOSA-N 0.000 claims description 3
- SBYVDRJAXWSXQL-AVGNSLFASA-N Glu-Asn-Phe Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O SBYVDRJAXWSXQL-AVGNSLFASA-N 0.000 claims description 3
- JRCUFCXYZLPSDZ-ACZMJKKPSA-N Glu-Asp-Ser Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(O)=O JRCUFCXYZLPSDZ-ACZMJKKPSA-N 0.000 claims description 3
- ISXJHXGYMJKXOI-GUBZILKMSA-N Glu-Cys-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CS)NC(=O)[C@@H](N)CCC(O)=O ISXJHXGYMJKXOI-GUBZILKMSA-N 0.000 claims description 3
- 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 claims description 3
- VFZIDQZAEBORGY-GLLZPBPUSA-N Glu-Gln-Thr Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O VFZIDQZAEBORGY-GLLZPBPUSA-N 0.000 claims description 3
- NKLRYVLERDYDBI-FXQIFTODSA-N Glu-Glu-Asp Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(O)=O)C(O)=O NKLRYVLERDYDBI-FXQIFTODSA-N 0.000 claims description 3
- BUZMZDDKFCSKOT-CIUDSAMLSA-N Glu-Glu-Glu Chemical compound OC(=O)CC[C@H](N)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O BUZMZDDKFCSKOT-CIUDSAMLSA-N 0.000 claims description 3
- IQACOVZVOMVILH-FXQIFTODSA-N Glu-Glu-Ser Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O IQACOVZVOMVILH-FXQIFTODSA-N 0.000 claims description 3
- KRGZZKWSBGPLKL-IUCAKERBSA-N Glu-Gly-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)CNC(=O)[C@H](CCC(=O)O)N KRGZZKWSBGPLKL-IUCAKERBSA-N 0.000 claims description 3
- ZWABFSSWTSAMQN-KBIXCLLPSA-N Glu-Ile-Ala Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](C)C(O)=O ZWABFSSWTSAMQN-KBIXCLLPSA-N 0.000 claims description 3
- IRXNJYPKBVERCW-DCAQKATOSA-N Glu-Leu-Glu Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(O)=O IRXNJYPKBVERCW-DCAQKATOSA-N 0.000 claims description 3
- QDMVXRNLOPTPIE-WDCWCFNPSA-N Glu-Lys-Thr Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)O)C(O)=O QDMVXRNLOPTPIE-WDCWCFNPSA-N 0.000 claims description 3
- XEKAJTCACGEBOK-KKUMJFAQSA-N Glu-Met-Phe Chemical compound CSCC[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H](CCC(=O)O)N XEKAJTCACGEBOK-KKUMJFAQSA-N 0.000 claims description 3
- QOXDAWODGSIDDI-GUBZILKMSA-N Glu-Ser-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CCC(=O)O)N QOXDAWODGSIDDI-GUBZILKMSA-N 0.000 claims description 3
- DTLLNDVORUEOTM-WDCWCFNPSA-N Glu-Thr-Lys Chemical compound [H]N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCCN)C(O)=O DTLLNDVORUEOTM-WDCWCFNPSA-N 0.000 claims description 3
- UZWUBBRJWFTHTD-LAEOZQHASA-N Glu-Val-Asn Chemical compound NC(=O)C[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CCC(O)=O UZWUBBRJWFTHTD-LAEOZQHASA-N 0.000 claims description 3
- SOYWRINXUSUWEQ-DLOVCJGASA-N Glu-Val-Val Chemical compound CC(C)[C@@H](C(O)=O)NC(=O)[C@H](C(C)C)NC(=O)[C@@H](N)CCC(O)=O SOYWRINXUSUWEQ-DLOVCJGASA-N 0.000 claims description 3
- SOEATRRYCIPEHA-BQBZGAKWSA-N Gly-Glu-Glu Chemical compound [H]NCC(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O SOEATRRYCIPEHA-BQBZGAKWSA-N 0.000 claims description 3
- STVHDEHTKFXBJQ-LAEOZQHASA-N Gly-Glu-Ile Chemical compound [H]NCC(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)CC)C(O)=O STVHDEHTKFXBJQ-LAEOZQHASA-N 0.000 claims description 3
- MBOAPAXLTUSMQI-JHEQGTHGSA-N Gly-Glu-Thr Chemical compound [H]NCC(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O MBOAPAXLTUSMQI-JHEQGTHGSA-N 0.000 claims description 3
- CCQOOWAONKGYKQ-BYPYZUCNSA-N Gly-Gly-Ala Chemical compound OC(=O)[C@H](C)NC(=O)CNC(=O)CN CCQOOWAONKGYKQ-BYPYZUCNSA-N 0.000 claims description 3
- AYBKPDHHVADEDA-YUMQZZPRSA-N Gly-His-Asn Chemical compound [H]NCC(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC(N)=O)C(O)=O AYBKPDHHVADEDA-YUMQZZPRSA-N 0.000 claims description 3
- PDUHNKAFQXQNLH-ZETCQYMHSA-N Gly-Lys-Gly Chemical compound NCCCC[C@H](NC(=O)CN)C(=O)NCC(O)=O PDUHNKAFQXQNLH-ZETCQYMHSA-N 0.000 claims description 3
- OCPPBNKYGYSLOE-IUCAKERBSA-N Gly-Pro-Met Chemical compound CSCC[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)CN OCPPBNKYGYSLOE-IUCAKERBSA-N 0.000 claims description 3
- SOEGEPHNZOISMT-BYPYZUCNSA-N Gly-Ser-Gly Chemical compound NCC(=O)N[C@@H](CO)C(=O)NCC(O)=O SOEGEPHNZOISMT-BYPYZUCNSA-N 0.000 claims description 3
- POJJAZJHBGXEGM-YUMQZZPRSA-N Gly-Ser-Lys Chemical compound C(CCN)C[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)CN POJJAZJHBGXEGM-YUMQZZPRSA-N 0.000 claims description 3
- RVKIPWVMZANZLI-UHFFFAOYSA-N H-Lys-Trp-OH Natural products C1=CC=C2C(CC(NC(=O)C(N)CCCCN)C(O)=O)=CNC2=C1 RVKIPWVMZANZLI-UHFFFAOYSA-N 0.000 claims description 3
- MWWOPNQSBXEUHO-ULQDDVLXSA-N His-Arg-Tyr Chemical compound C([C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CC=1C=CC(O)=CC=1)C(O)=O)C1=CN=CN1 MWWOPNQSBXEUHO-ULQDDVLXSA-N 0.000 claims description 3
- IDQKGZWUPVOGPZ-GUBZILKMSA-N His-Cys-Gln Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CS)C(=O)N[C@@H](CCC(=O)N)C(=O)O)N IDQKGZWUPVOGPZ-GUBZILKMSA-N 0.000 claims description 3
- HIAHVKLTHNOENC-HGNGGELXSA-N His-Glu-Ala Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O HIAHVKLTHNOENC-HGNGGELXSA-N 0.000 claims description 3
- PQKCQZHAGILVIM-NKIYYHGXSA-N His-Glu-Thr Chemical compound C[C@@H](O)[C@H](NC(=O)[C@H](CCC(O)=O)NC(=O)[C@@H](N)Cc1cnc[nH]1)C(O)=O PQKCQZHAGILVIM-NKIYYHGXSA-N 0.000 claims description 3
- ZHHLTWUOWXHVQJ-YUMQZZPRSA-N His-Ser-Gly Chemical compound C1=C(NC=N1)C[C@@H](C(=O)N[C@@H](CO)C(=O)NCC(=O)O)N ZHHLTWUOWXHVQJ-YUMQZZPRSA-N 0.000 claims description 3
- PZAJPILZRFPYJJ-SRVKXCTJSA-N His-Ser-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(O)=O PZAJPILZRFPYJJ-SRVKXCTJSA-N 0.000 claims description 3
- VIJMRAIWYWRXSR-CIUDSAMLSA-N His-Ser-Ser Chemical compound OC[C@@H](C(O)=O)NC(=O)[C@H](CO)NC(=O)[C@@H](N)CC1=CN=CN1 VIJMRAIWYWRXSR-CIUDSAMLSA-N 0.000 claims description 3
- LNVILFYCPVOHPV-IHPCNDPISA-N His-Trp-Leu Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CC(C)C)C(O)=O LNVILFYCPVOHPV-IHPCNDPISA-N 0.000 claims description 3
- WSWAUVHXQREQQG-JYJNAYRXSA-N His-Tyr-Gln Chemical compound [H]N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O WSWAUVHXQREQQG-JYJNAYRXSA-N 0.000 claims description 3
- DFJJAVZIHDFOGQ-MNXVOIDGSA-N Ile-Glu-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CCCCN)C(=O)O)N DFJJAVZIHDFOGQ-MNXVOIDGSA-N 0.000 claims description 3
- TVYWVSJGSHQWMT-AJNGGQMLSA-N Ile-Leu-Lys Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)O)N TVYWVSJGSHQWMT-AJNGGQMLSA-N 0.000 claims description 3
- JHNJNTMTZHEDLJ-NAKRPEOUSA-N Ile-Ser-Arg Chemical compound CC[C@H](C)[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O JHNJNTMTZHEDLJ-NAKRPEOUSA-N 0.000 claims description 3
- PELCGFMHLZXWBQ-BJDJZHNGSA-N Ile-Ser-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CC(C)C)C(=O)O)N PELCGFMHLZXWBQ-BJDJZHNGSA-N 0.000 claims description 3
- 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 claims description 3
- KBDIBHQICWDGDL-PPCPHDFISA-N Ile-Thr-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(C)C)C(=O)O)N KBDIBHQICWDGDL-PPCPHDFISA-N 0.000 claims description 3
- HASRFYOMVPJRPU-SRVKXCTJSA-N Leu-Arg-Glu Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCCN=C(N)N)C(=O)N[C@@H](CCC(O)=O)C(O)=O HASRFYOMVPJRPU-SRVKXCTJSA-N 0.000 claims description 3
- JQSXWJXBASFONF-KKUMJFAQSA-N Leu-Asp-Phe Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O JQSXWJXBASFONF-KKUMJFAQSA-N 0.000 claims description 3
- PVMPDMIKUVNOBD-CIUDSAMLSA-N Leu-Asp-Ser Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CO)C(O)=O PVMPDMIKUVNOBD-CIUDSAMLSA-N 0.000 claims description 3
- KAFOIVJDVSZUMD-DCAQKATOSA-N Leu-Gln-Gln Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCC(N)=O)C(O)=O KAFOIVJDVSZUMD-DCAQKATOSA-N 0.000 claims description 3
- KAFOIVJDVSZUMD-UHFFFAOYSA-N Leu-Gln-Gln Natural products CC(C)CC(N)C(=O)NC(CCC(N)=O)C(=O)NC(CCC(N)=O)C(O)=O KAFOIVJDVSZUMD-UHFFFAOYSA-N 0.000 claims description 3
- FQZPTCNSNPWHLJ-AVGNSLFASA-N Leu-Gln-Lys Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCCCN)C(O)=O FQZPTCNSNPWHLJ-AVGNSLFASA-N 0.000 claims description 3
- CIVKXGPFXDIQBV-WDCWCFNPSA-N Leu-Gln-Thr Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H]([C@@H](C)O)C(O)=O CIVKXGPFXDIQBV-WDCWCFNPSA-N 0.000 claims description 3
- DZQMXBALGUHGJT-GUBZILKMSA-N Leu-Glu-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](C)C(O)=O DZQMXBALGUHGJT-GUBZILKMSA-N 0.000 claims description 3
- OGUUKPXUTHOIAV-SDDRHHMPSA-N Leu-Glu-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N OGUUKPXUTHOIAV-SDDRHHMPSA-N 0.000 claims description 3
- APFJUBGRZGMQFF-QWRGUYRKSA-N Leu-Gly-Lys Chemical compound CC(C)C[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCCCN APFJUBGRZGMQFF-QWRGUYRKSA-N 0.000 claims description 3
- XBCWOTOCBXXJDG-BZSNNMDCSA-N Leu-His-Phe Chemical compound C([C@H](NC(=O)[C@@H](N)CC(C)C)C(=O)N[C@@H](CC=1C=CC=CC=1)C(O)=O)C1=CN=CN1 XBCWOTOCBXXJDG-BZSNNMDCSA-N 0.000 claims description 3
- ZRHDPZAAWLXXIR-SRVKXCTJSA-N Leu-Lys-Ala Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](C)C(O)=O ZRHDPZAAWLXXIR-SRVKXCTJSA-N 0.000 claims description 3
- LVTJJOJKDCVZGP-QWRGUYRKSA-N Leu-Lys-Gly Chemical compound [H]N[C@@H](CC(C)C)C(=O)N[C@@H](CCCCN)C(=O)NCC(O)=O LVTJJOJKDCVZGP-QWRGUYRKSA-N 0.000 claims description 3
- PKKMDPNFGULLNQ-AVGNSLFASA-N Leu-Met-Arg Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCCN=C(N)N)C(O)=O PKKMDPNFGULLNQ-AVGNSLFASA-N 0.000 claims description 3
- AMSSKPUHBUQBOQ-SRVKXCTJSA-N Leu-Ser-Lys Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CO)C(=O)N[C@@H](CCCCN)C(=O)O)N AMSSKPUHBUQBOQ-SRVKXCTJSA-N 0.000 claims description 3
- IWMJFLJQHIDZQW-KKUMJFAQSA-N Leu-Ser-Phe Chemical compound CC(C)C[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 IWMJFLJQHIDZQW-KKUMJFAQSA-N 0.000 claims description 3
- SBANPBVRHYIMRR-GARJFASQSA-N Leu-Ser-Pro Chemical compound CC(C)C[C@@H](C(=O)N[C@@H](CO)C(=O)N1CCC[C@@H]1C(=O)O)N SBANPBVRHYIMRR-GARJFASQSA-N 0.000 claims description 3
- SBANPBVRHYIMRR-UHFFFAOYSA-N Leu-Ser-Pro Natural products CC(C)CC(N)C(=O)NC(CO)C(=O)N1CCCC1C(O)=O SBANPBVRHYIMRR-UHFFFAOYSA-N 0.000 claims description 3
- NQCJGQHHYZNUDK-DCAQKATOSA-N Lys-Arg-Ser Chemical compound NCCCC[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](CO)C(O)=O)CCCN=C(N)N NQCJGQHHYZNUDK-DCAQKATOSA-N 0.000 claims description 3
- KSFQPRLZAUXXPT-GARJFASQSA-N Lys-Cys-Pro Chemical compound C1C[C@@H](N(C1)C(=O)[C@H](CS)NC(=O)[C@H](CCCCN)N)C(=O)O KSFQPRLZAUXXPT-GARJFASQSA-N 0.000 claims description 3
- WTZUSCUIVPVCRH-SRVKXCTJSA-N Lys-Gln-Arg Chemical compound NCCCC[C@H](N)C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@H](C(O)=O)CCCN=C(N)N WTZUSCUIVPVCRH-SRVKXCTJSA-N 0.000 claims description 3
- NNCDAORZCMPZPX-GUBZILKMSA-N Lys-Gln-Ser Chemical compound C(CCN)C[C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H](CO)C(=O)O)N NNCDAORZCMPZPX-GUBZILKMSA-N 0.000 claims description 3
- PBIPLDMFHAICIP-DCAQKATOSA-N Lys-Glu-Glu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O PBIPLDMFHAICIP-DCAQKATOSA-N 0.000 claims description 3
- DUTMKEAPLLUGNO-JYJNAYRXSA-N Lys-Glu-Phe Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O DUTMKEAPLLUGNO-JYJNAYRXSA-N 0.000 claims description 3
- VEGLGAOVLFODGC-GUBZILKMSA-N Lys-Glu-Ser Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CO)C(O)=O VEGLGAOVLFODGC-GUBZILKMSA-N 0.000 claims description 3
- PGLGNCVOWIORQE-SRVKXCTJSA-N Lys-His-Ser Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC1=CNC=N1)C(=O)N[C@@H](CO)C(O)=O PGLGNCVOWIORQE-SRVKXCTJSA-N 0.000 claims description 3
- NCZIQZYZPUPMKY-PPCPHDFISA-N Lys-Ile-Thr Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H]([C@@H](C)O)C(O)=O NCZIQZYZPUPMKY-PPCPHDFISA-N 0.000 claims description 3
- MYZMQWHPDAYKIE-SRVKXCTJSA-N Lys-Leu-Ala Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](C)C(O)=O MYZMQWHPDAYKIE-SRVKXCTJSA-N 0.000 claims description 3
- YUAXTFMFMOIMAM-QWRGUYRKSA-N Lys-Lys-Gly Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCCCN)C(=O)NCC(O)=O YUAXTFMFMOIMAM-QWRGUYRKSA-N 0.000 claims description 3
- ZCWWVXAXWUAEPZ-SRVKXCTJSA-N Lys-Met-Glu Chemical compound [H]N[C@@H](CCCCN)C(=O)N[C@@H](CCSC)C(=O)N[C@@H](CCC(O)=O)C(O)=O ZCWWVXAXWUAEPZ-SRVKXCTJSA-N 0.000 claims description 3
- KFSALEZVQJYHCE-AVGNSLFASA-N Lys-Met-Val Chemical compound CC(C)[C@@H](C(=O)O)NC(=O)[C@H](CCSC)NC(=O)[C@H](CCCCN)N KFSALEZVQJYHCE-AVGNSLFASA-N 0.000 claims description 3
- 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 claims description 3
- DYJOORGDQIGZAS-DCAQKATOSA-N Lys-Ser-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@H](CO)NC(=O)[C@H](CCCCN)N DYJOORGDQIGZAS-DCAQKATOSA-N 0.000 claims description 3
- KXYLFJIQDIMURW-IHPCNDPISA-N Lys-Trp-Leu Chemical compound C1=CC=C2C(C[C@@H](C(=O)N[C@@H](CC(C)C)C(O)=O)NC(=O)[C@@H](N)CCCCN)=CNC2=C1 KXYLFJIQDIMURW-IHPCNDPISA-N 0.000 claims description 3
- 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 claims description 3
- BLIPQDLSCFGUFA-GUBZILKMSA-N Met-Arg-Asn Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CC(N)=O)C(O)=O BLIPQDLSCFGUFA-GUBZILKMSA-N 0.000 claims description 3
- XOMXAVJBLRROMC-IHRRRGAJSA-N Met-Asp-Phe Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@H](C(O)=O)CC1=CC=CC=C1 XOMXAVJBLRROMC-IHRRRGAJSA-N 0.000 claims description 3
- VOOINLQYUZOREH-SRVKXCTJSA-N Met-Gln-Leu Chemical compound CC(C)C[C@@H](C(=O)O)NC(=O)[C@H](CCC(=O)N)NC(=O)[C@H](CCSC)N VOOINLQYUZOREH-SRVKXCTJSA-N 0.000 claims description 3
- BMHIFARYXOJDLD-WPRPVWTQSA-N Met-Gly-Val Chemical compound [H]N[C@@H](CCSC)C(=O)NCC(=O)N[C@@H](C(C)C)C(O)=O BMHIFARYXOJDLD-WPRPVWTQSA-N 0.000 claims description 3
- NHXXGBXJTLRGJI-GUBZILKMSA-N Met-Pro-Ser Chemical compound [H]N[C@@H](CCSC)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CO)C(O)=O NHXXGBXJTLRGJI-GUBZILKMSA-N 0.000 claims description 3
- HLZORBMOISUNIV-DCAQKATOSA-N Met-Ser-Leu Chemical compound CSCC[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CC(C)C HLZORBMOISUNIV-DCAQKATOSA-N 0.000 claims description 3
- PHURAEXVWLDIGT-LPEHRKFASA-N Met-Ser-Pro Chemical compound CSCC[C@@H](C(=O)N[C@@H](CO)C(=O)N1CCC[C@@H]1C(=O)O)N PHURAEXVWLDIGT-LPEHRKFASA-N 0.000 claims description 3
- 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 claims description 3
- XZFYRXDAULDNFX-UHFFFAOYSA-N N-L-cysteinyl-L-phenylalanine Natural products SCC(N)C(=O)NC(C(O)=O)CC1=CC=CC=C1 XZFYRXDAULDNFX-UHFFFAOYSA-N 0.000 claims description 3
- 108010002311 N-glycylglutamic acid Proteins 0.000 claims description 3
- 108010087066 N2-tryptophyllysine Proteins 0.000 claims description 3
- BQVUABVGYYSDCJ-UHFFFAOYSA-N Nalpha-L-Leucyl-L-tryptophan Natural products C1=CC=C2C(CC(NC(=O)C(N)CC(C)C)C(O)=O)=CNC2=C1 BQVUABVGYYSDCJ-UHFFFAOYSA-N 0.000 claims description 3
- KXUZHWXENMYOHC-QEJZJMRPSA-N Phe-Leu-Ala Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC(C)C)C(=O)N[C@@H](C)C(O)=O KXUZHWXENMYOHC-QEJZJMRPSA-N 0.000 claims description 3
- KAJLHCWRWDSROH-BZSNNMDCSA-N Phe-Phe-Asp Chemical compound C([C@H](N)C(=O)N[C@@H](CC=1C=CC=CC=1)C(=O)N[C@@H](CC(O)=O)C(O)=O)C1=CC=CC=C1 KAJLHCWRWDSROH-BZSNNMDCSA-N 0.000 claims description 3
- DSXPMZMSJHOKKK-HJOGWXRNSA-N Phe-Phe-Tyr Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O DSXPMZMSJHOKKK-HJOGWXRNSA-N 0.000 claims description 3
- MCIXMYKSPQUMJG-SRVKXCTJSA-N Phe-Ser-Ser Chemical compound [H]N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CO)C(=O)N[C@@H](CO)C(O)=O MCIXMYKSPQUMJG-SRVKXCTJSA-N 0.000 claims description 3
- DRIJZWBRGMJCDD-DCAQKATOSA-N Pro-Gln-Met Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CCC(N)=O)C(=O)N[C@@H](CCSC)C(O)=O DRIJZWBRGMJCDD-DCAQKATOSA-N 0.000 claims description 3
- BUEIYHBJHCDAMI-UFYCRDLUSA-N Pro-Phe-Phe Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CC1=CC=CC=C1)C(O)=O BUEIYHBJHCDAMI-UFYCRDLUSA-N 0.000 claims description 3
- QUBVFEANYYWBTM-VEVYYDQMSA-N Pro-Thr-Asp Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(O)=O QUBVFEANYYWBTM-VEVYYDQMSA-N 0.000 claims description 3
- IURWWZYKYPEANQ-HJGDQZAQSA-N Pro-Thr-Glu Chemical compound [H]N1CCC[C@H]1C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCC(O)=O)C(O)=O IURWWZYKYPEANQ-HJGDQZAQSA-N 0.000 claims description 3
- YUSRGTQIPCJNHQ-CIUDSAMLSA-N Ser-Arg-Glu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCC(O)=O)C(O)=O YUSRGTQIPCJNHQ-CIUDSAMLSA-N 0.000 claims description 3
- DBIDZNUXSLXVRG-FXQIFTODSA-N Ser-Asp-Met Chemical compound CSCC[C@@H](C(=O)O)NC(=O)[C@H](CC(=O)O)NC(=O)[C@H](CO)N DBIDZNUXSLXVRG-FXQIFTODSA-N 0.000 claims description 3
- LALNXSXEYFUUDD-GUBZILKMSA-N Ser-Glu-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CCC(O)=O)C(=O)N[C@@H](CC(C)C)C(O)=O LALNXSXEYFUUDD-GUBZILKMSA-N 0.000 claims description 3
- MUARUIBTKQJKFY-WHFBIAKZSA-N Ser-Gly-Asp Chemical compound [H]N[C@@H](CO)C(=O)NCC(=O)N[C@@H](CC(O)=O)C(O)=O MUARUIBTKQJKFY-WHFBIAKZSA-N 0.000 claims description 3
- MIJWOJAXARLEHA-WDSKDSINSA-N Ser-Gly-Glu Chemical compound OC[C@H](N)C(=O)NCC(=O)N[C@H](C(O)=O)CCC(O)=O MIJWOJAXARLEHA-WDSKDSINSA-N 0.000 claims description 3
- UIPXCLNLUUAMJU-JBDRJPRFSA-N Ser-Ile-Ser Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CO)C(O)=O UIPXCLNLUUAMJU-JBDRJPRFSA-N 0.000 claims description 3
- MQQBBLVOUUJKLH-HJPIBITLSA-N Ser-Ile-Tyr Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H]([C@@H](C)CC)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(O)=O MQQBBLVOUUJKLH-HJPIBITLSA-N 0.000 claims description 3
- CRJZZXMAADSBBQ-SRVKXCTJSA-N Ser-Lys-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@H](CCCCN)NC(=O)[C@@H](N)CO CRJZZXMAADSBBQ-SRVKXCTJSA-N 0.000 claims description 3
- JAWGSPUJAXYXJA-IHRRRGAJSA-N Ser-Phe-Arg Chemical compound NC(N)=NCCC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@H](CO)N)CC1=CC=CC=C1 JAWGSPUJAXYXJA-IHRRRGAJSA-N 0.000 claims description 3
- FZEUTKVQGMVGHW-AVGNSLFASA-N Ser-Phe-Gln Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O FZEUTKVQGMVGHW-AVGNSLFASA-N 0.000 claims description 3
- XKFJENWJGHMDLI-QWRGUYRKSA-N Ser-Phe-Gly Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)NCC(O)=O XKFJENWJGHMDLI-QWRGUYRKSA-N 0.000 claims description 3
- NUEHQDHDLDXCRU-GUBZILKMSA-N Ser-Pro-Arg Chemical compound OC[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCCN=C(N)N)C(O)=O NUEHQDHDLDXCRU-GUBZILKMSA-N 0.000 claims description 3
- FKYWFUYPVKLJLP-DCAQKATOSA-N Ser-Pro-Leu Chemical compound CC(C)C[C@@H](C(O)=O)NC(=O)[C@@H]1CCCN1C(=O)[C@@H](N)CO FKYWFUYPVKLJLP-DCAQKATOSA-N 0.000 claims description 3
- FZXOPYUEQGDGMS-ACZMJKKPSA-N Ser-Ser-Gln Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](CO)C(=O)N[C@@H](CCC(N)=O)C(O)=O FZXOPYUEQGDGMS-ACZMJKKPSA-N 0.000 claims description 3
- QNBVFKZSSRYNFX-CUJWVEQBSA-N Ser-Thr-His Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CC1=CN=CN1)C(=O)O)NC(=O)[C@H](CO)N)O QNBVFKZSSRYNFX-CUJWVEQBSA-N 0.000 claims description 3
- PCJLFYBAQZQOFE-KATARQTJSA-N Ser-Thr-Lys Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CCCCN)C(=O)O)NC(=O)[C@H](CO)N)O PCJLFYBAQZQOFE-KATARQTJSA-N 0.000 claims description 3
- YEDSOSIKVUMIJE-DCAQKATOSA-N Ser-Val-Leu Chemical compound [H]N[C@@H](CO)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](CC(C)C)C(O)=O YEDSOSIKVUMIJE-DCAQKATOSA-N 0.000 claims description 3
- CAGTXGDOIFXLPC-KZVJFYERSA-N Thr-Arg-Ala Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@H](C(=O)N[C@@H](C)C(O)=O)CCCN=C(N)N CAGTXGDOIFXLPC-KZVJFYERSA-N 0.000 claims description 3
- XYEXCEPTALHNEV-RCWTZXSCSA-N Thr-Arg-Arg Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CCCNC(N)=N)C(=O)N[C@@H](CCCNC(N)=N)C(O)=O XYEXCEPTALHNEV-RCWTZXSCSA-N 0.000 claims description 3
- OJRNZRROAIAHDL-LKXGYXEUSA-N Thr-Asn-Ser Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(N)=O)C(=O)N[C@@H](CO)C(O)=O OJRNZRROAIAHDL-LKXGYXEUSA-N 0.000 claims description 3
- YBXMGKCLOPDEKA-NUMRIWBASA-N Thr-Asp-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC(O)=O)C(=O)N[C@@H](CCC(O)=O)C(O)=O YBXMGKCLOPDEKA-NUMRIWBASA-N 0.000 claims description 3
- KGKWKSSSQGGYAU-SUSMZKCASA-N Thr-Gln-Thr Chemical compound C[C@H]([C@@H](C(=O)N[C@@H](CCC(=O)N)C(=O)N[C@@H]([C@@H](C)O)C(=O)O)N)O KGKWKSSSQGGYAU-SUSMZKCASA-N 0.000 claims description 3
- ADPHPKGWVDHWML-PPCPHDFISA-N Thr-Ile-Leu Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC(C)C)C(=O)O)NC(=O)[C@H]([C@@H](C)O)N ADPHPKGWVDHWML-PPCPHDFISA-N 0.000 claims description 3
- PZSDPRBZINDEJV-HTUGSXCWSA-N Thr-Phe-Gln Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)N[C@@H](CCC(N)=O)C(O)=O PZSDPRBZINDEJV-HTUGSXCWSA-N 0.000 claims description 3
- 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 claims description 3
- XKWABWFMQXMUMT-HJGDQZAQSA-N Thr-Pro-Glu Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCC(O)=O)C(O)=O XKWABWFMQXMUMT-HJGDQZAQSA-N 0.000 claims description 3
- GFRIEEKFXOVPIR-RHYQMDGZSA-N Thr-Pro-Lys Chemical compound C[C@@H](O)[C@H](N)C(=O)N1CCC[C@H]1C(=O)N[C@@H](CCCCN)C(O)=O GFRIEEKFXOVPIR-RHYQMDGZSA-N 0.000 claims description 3
- IQPWNQRRAJHOKV-KATARQTJSA-N Thr-Ser-Lys Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCCCN IQPWNQRRAJHOKV-KATARQTJSA-N 0.000 claims description 3
- UQCNIMDPYICBTR-KYNKHSRBSA-N Thr-Thr-Gly Chemical compound [H]N[C@@H]([C@@H](C)O)C(=O)N[C@@H]([C@@H](C)O)C(=O)NCC(O)=O UQCNIMDPYICBTR-KYNKHSRBSA-N 0.000 claims description 3
- OGOYMQWIWHGTGH-KZVJFYERSA-N Thr-Val-Ala Chemical compound C[C@@H](O)[C@H](N)C(=O)N[C@@H](C(C)C)C(=O)N[C@@H](C)C(O)=O OGOYMQWIWHGTGH-KZVJFYERSA-N 0.000 claims description 3
- SCQBNMKLZVCXNX-ZFWWWQNUSA-N Trp-Arg-Gly Chemical compound C1=CC=C2C(=C1)C(=CN2)C[C@@H](C(=O)N[C@@H](CCCN=C(N)N)C(=O)NCC(=O)O)N SCQBNMKLZVCXNX-ZFWWWQNUSA-N 0.000 claims description 3
- BYSKNUASOAGJSS-NQCBNZPSSA-N Trp-Ile-Phe Chemical compound CC[C@H](C)[C@@H](C(=O)N[C@@H](CC1=CC=CC=C1)C(=O)O)NC(=O)[C@H](CC2=CNC3=CC=CC=C32)N BYSKNUASOAGJSS-NQCBNZPSSA-N 0.000 claims description 3
- NWQCKAPDGQMZQN-IHPCNDPISA-N Trp-Lys-Leu Chemical compound [H]N[C@@H](CC1=CNC2=C1C=CC=C2)C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(C)C)C(O)=O NWQCKAPDGQMZQN-IHPCNDPISA-N 0.000 claims description 3
- BXJQKVDPRMLGKN-PMVMPFDFSA-N Tyr-Trp-Leu Chemical compound C([C@H](N)C(=O)N[C@@H](CC=1C2=CC=CC=C2NC=1)C(=O)N[C@@H](CC(C)C)C(O)=O)C1=CC=C(O)C=C1 BXJQKVDPRMLGKN-PMVMPFDFSA-N 0.000 claims description 3
- GZWPQZDVTBZVEP-BZSNNMDCSA-N Tyr-Tyr-Asn Chemical compound [H]N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC1=CC=C(O)C=C1)C(=O)N[C@@H](CC(N)=O)C(O)=O GZWPQZDVTBZVEP-BZSNNMDCSA-N 0.000 claims description 3
- DDNIHOWRDOXXPF-NGZCFLSTSA-N Val-Asp-Pro Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC(=O)O)C(=O)N1CCC[C@@H]1C(=O)O)N DDNIHOWRDOXXPF-NGZCFLSTSA-N 0.000 claims description 3
- ZXAGTABZUOMUDO-GVXVVHGQSA-N Val-Glu-Lys Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCC(=O)O)C(=O)N[C@@H](CCCCN)C(=O)O)N ZXAGTABZUOMUDO-GVXVVHGQSA-N 0.000 claims description 3
- SYOMXKPPFZRELL-ONGXEEELSA-N Val-Gly-Lys Chemical compound CC(C)[C@@H](C(=O)NCC(=O)N[C@@H](CCCCN)C(=O)O)N SYOMXKPPFZRELL-ONGXEEELSA-N 0.000 claims description 3
- 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 claims description 3
- WDIWOIRFNMLNKO-ULQDDVLXSA-N Val-Leu-Tyr Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](CC(C)C)C(=O)N[C@H](C(O)=O)CC1=CC=C(O)C=C1 WDIWOIRFNMLNKO-ULQDDVLXSA-N 0.000 claims description 3
- KTEZUXISLQTDDQ-NHCYSSNCSA-N Val-Lys-Asp Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CCCCN)C(=O)N[C@@H](CC(=O)O)C(=O)O)N KTEZUXISLQTDDQ-NHCYSSNCSA-N 0.000 claims description 3
- VIKZGAUAKQZDOF-NRPADANISA-N Val-Ser-Glu Chemical compound CC(C)[C@H](N)C(=O)N[C@@H](CO)C(=O)N[C@H](C(O)=O)CCC(O)=O VIKZGAUAKQZDOF-NRPADANISA-N 0.000 claims description 3
- HTONZBWRYUKUKC-RCWTZXSCSA-N Val-Thr-Val Chemical compound CC(C)[C@H](N)C(=O)N[C@@H]([C@@H](C)O)C(=O)N[C@@H](C(C)C)C(O)=O HTONZBWRYUKUKC-RCWTZXSCSA-N 0.000 claims description 3
- AYHNXCJKBLYVOA-KSZLIROESA-N Val-Trp-Pro Chemical compound CC(C)[C@@H](C(=O)N[C@@H](CC1=CNC2=CC=CC=C21)C(=O)N3CCC[C@@H]3C(=O)O)N AYHNXCJKBLYVOA-KSZLIROESA-N 0.000 claims description 3
- RLVTVHSDKHBFQP-ULQDDVLXSA-N Val-Tyr-Lys Chemical compound NCCCC[C@@H](C(O)=O)NC(=O)[C@@H](NC(=O)[C@@H](N)C(C)C)CC1=CC=C(O)C=C1 RLVTVHSDKHBFQP-ULQDDVLXSA-N 0.000 claims description 3
- 108010076324 alanyl-glycyl-glycine Proteins 0.000 claims description 3
- 108010009111 arginyl-glycyl-glutamic acid Proteins 0.000 claims description 3
- 108010040443 aspartyl-aspartic acid Proteins 0.000 claims description 3
- 108010038633 aspartylglutamate Proteins 0.000 claims description 3
- 108010063718 gamma-glutamylaspartic acid Proteins 0.000 claims description 3
- 108010042598 glutamyl-aspartyl-glycine Proteins 0.000 claims description 3
- VPZXBVLAVMBEQI-UHFFFAOYSA-N glycyl-DL-alpha-alanine Natural products OC(=O)C(C)NC(=O)CN VPZXBVLAVMBEQI-UHFFFAOYSA-N 0.000 claims description 3
- JYPCXBJRLBHWME-UHFFFAOYSA-N glycyl-L-prolyl-L-arginine Natural products NCC(=O)N1CCCC1C(=O)NC(CCCN=C(N)N)C(O)=O JYPCXBJRLBHWME-UHFFFAOYSA-N 0.000 claims description 3
- 108010020688 glycylhistidine Proteins 0.000 claims description 3
- 108010015792 glycyllysine Proteins 0.000 claims description 3
- 108010057821 leucylproline Proteins 0.000 claims description 3
- 108010054155 lysyllysine Proteins 0.000 claims description 3
- 108010038320 lysylphenylalanine Proteins 0.000 claims description 3
- 108010084572 phenylalanyl-valine Proteins 0.000 claims description 3
- 108010031719 prolyl-serine Proteins 0.000 claims description 3
- 108010071207 serylmethionine Proteins 0.000 claims description 3
- 230000001629 suppression Effects 0.000 claims description 3
- 108010031491 threonyl-lysyl-glutamic acid Proteins 0.000 claims description 3
- 108010072986 threonyl-seryl-lysine Proteins 0.000 claims description 3
- 108010015666 tryptophyl-leucyl-glutamic acid Proteins 0.000 claims description 3
- 108010045269 tryptophyltryptophan Proteins 0.000 claims description 3
- 108010051110 tyrosyl-lysine Proteins 0.000 claims description 3
- 230000002018 overexpression Effects 0.000 claims description 2
- 150000001413 amino acids Chemical group 0.000 claims 1
- 108091008324 binding proteins Proteins 0.000 claims 1
- 230000003111 delayed effect Effects 0.000 claims 1
- 102000004169 proteins and genes Human genes 0.000 abstract description 17
- 238000003780 insertion Methods 0.000 abstract description 9
- 230000037431 insertion Effects 0.000 abstract description 9
- 125000003275 alpha amino acid group Chemical group 0.000 abstract description 6
- 230000000694 effects Effects 0.000 abstract description 5
- 238000012271 agricultural production Methods 0.000 abstract description 2
- 125000000539 amino acid group Chemical group 0.000 abstract description 2
- 108091029865 Exogenous DNA Proteins 0.000 abstract 1
- 230000037430 deletion Effects 0.000 abstract 1
- 238000012217 deletion Methods 0.000 abstract 1
- 238000006467 substitution reaction Methods 0.000 abstract 1
- 241000196324 Embryophyta Species 0.000 description 32
- 108020004414 DNA Proteins 0.000 description 19
- 210000004027 cell Anatomy 0.000 description 14
- 235000018102 proteins Nutrition 0.000 description 11
- 238000012163 sequencing technique Methods 0.000 description 10
- 238000010378 bimolecular fluorescence complementation Methods 0.000 description 9
- 238000000034 method Methods 0.000 description 9
- 238000004458 analytical method Methods 0.000 description 8
- 230000001276 controlling effect Effects 0.000 description 8
- 108091008146 restriction endonucleases Proteins 0.000 description 8
- 238000003757 reverse transcription PCR Methods 0.000 description 8
- 229910001424 calcium ion Inorganic materials 0.000 description 7
- 238000002474 experimental method Methods 0.000 description 7
- 239000013642 negative control Substances 0.000 description 7
- 239000013612 plasmid Substances 0.000 description 7
- 238000003158 yeast two-hybrid assay Methods 0.000 description 7
- BHPQYMZQTOCNFJ-UHFFFAOYSA-N Calcium cation Chemical compound [Ca+2] BHPQYMZQTOCNFJ-UHFFFAOYSA-N 0.000 description 6
- 101000702488 Rattus norvegicus High affinity cationic amino acid transporter 1 Proteins 0.000 description 6
- 238000013459 approach Methods 0.000 description 6
- 230000000295 complement effect Effects 0.000 description 6
- 230000012010 growth Effects 0.000 description 6
- 230000035772 mutation Effects 0.000 description 6
- 210000005253 yeast cell Anatomy 0.000 description 6
- 240000004808 Saccharomyces cerevisiae Species 0.000 description 5
- 238000005286 illumination Methods 0.000 description 5
- 230000008569 process Effects 0.000 description 5
- 238000011160 research Methods 0.000 description 5
- 210000000582 semen Anatomy 0.000 description 5
- 210000002966 serum Anatomy 0.000 description 5
- 244000291564 Allium cepa Species 0.000 description 4
- 235000002732 Allium cepa var. cepa Nutrition 0.000 description 4
- 241000219195 Arabidopsis thaliana Species 0.000 description 4
- 230000003321 amplification Effects 0.000 description 4
- 230000027455 binding Effects 0.000 description 4
- 238000010367 cloning Methods 0.000 description 4
- MURGITYSBWUQTI-UHFFFAOYSA-N fluorescin Chemical compound OC(=O)C1=CC=CC=C1C1C2=CC=C(O)C=C2OC2=CC(O)=CC=C21 MURGITYSBWUQTI-UHFFFAOYSA-N 0.000 description 4
- 239000007952 growth promoter Substances 0.000 description 4
- 230000006698 induction Effects 0.000 description 4
- 239000000463 material Substances 0.000 description 4
- 238000003199 nucleic acid amplification method Methods 0.000 description 4
- 108091032973 (ribonucleotides)n+m Proteins 0.000 description 3
- 101100397135 Arabidopsis thaliana IQM2 gene Proteins 0.000 description 3
- 241000628997 Flos Species 0.000 description 3
- 238000010240 RT-PCR analysis Methods 0.000 description 3
- 238000006243 chemical reaction Methods 0.000 description 3
- 210000002615 epidermis Anatomy 0.000 description 3
- 239000013604 expression vector Substances 0.000 description 3
- 230000002068 genetic effect Effects 0.000 description 3
- 230000007246 mechanism Effects 0.000 description 3
- 238000001086 yeast two-hybrid system Methods 0.000 description 3
- 108090000790 Enzymes Proteins 0.000 description 2
- 102000004190 Enzymes Human genes 0.000 description 2
- 108700039887 Essential Genes Proteins 0.000 description 2
- 102100039556 Galectin-4 Human genes 0.000 description 2
- 101000608765 Homo sapiens Galectin-4 Proteins 0.000 description 2
- 108090000862 Ion Channels Proteins 0.000 description 2
- 102000004310 Ion Channels Human genes 0.000 description 2
- 240000002853 Nelumbo nucifera Species 0.000 description 2
- 235000006508 Nelumbo nucifera Nutrition 0.000 description 2
- 235000006510 Nelumbo pentapetala Nutrition 0.000 description 2
- 244000061176 Nicotiana tabacum Species 0.000 description 2
- 235000002637 Nicotiana tabacum Nutrition 0.000 description 2
- 230000004913 activation Effects 0.000 description 2
- 239000011575 calcium Substances 0.000 description 2
- 230000008859 change Effects 0.000 description 2
- 238000010276 construction Methods 0.000 description 2
- 238000001514 detection method Methods 0.000 description 2
- 238000011161 development Methods 0.000 description 2
- 230000018109 developmental process Effects 0.000 description 2
- 235000013399 edible fruits Nutrition 0.000 description 2
- 238000005516 engineering process Methods 0.000 description 2
- 230000007613 environmental effect Effects 0.000 description 2
- 230000004927 fusion Effects 0.000 description 2
- 108020001507 fusion proteins Proteins 0.000 description 2
- 102000037865 fusion proteins Human genes 0.000 description 2
- 238000010353 genetic engineering Methods 0.000 description 2
- 239000001963 growth medium Substances 0.000 description 2
- 230000003993 interaction Effects 0.000 description 2
- 238000004519 manufacturing process Methods 0.000 description 2
- 239000002609 medium Substances 0.000 description 2
- 108020004999 messenger RNA Proteins 0.000 description 2
- 239000000203 mixture Substances 0.000 description 2
- 235000015097 nutrients Nutrition 0.000 description 2
- 235000016709 nutrition Nutrition 0.000 description 2
- 230000035764 nutrition Effects 0.000 description 2
- 239000002245 particle Substances 0.000 description 2
- 230000037361 pathway Effects 0.000 description 2
- 239000013641 positive control Substances 0.000 description 2
- 230000001105 regulatory effect Effects 0.000 description 2
- 239000000523 sample Substances 0.000 description 2
- 238000012216 screening Methods 0.000 description 2
- 230000019491 signal transduction Effects 0.000 description 2
- 239000002689 soil Substances 0.000 description 2
- 238000004659 sterilization and disinfection Methods 0.000 description 2
- XLYOFNOQVPJJNP-UHFFFAOYSA-N water Chemical compound O XLYOFNOQVPJJNP-UHFFFAOYSA-N 0.000 description 2
- IXVMHGVQKLDRKH-YEJCTVDLSA-N (22s,23s)-epibrassinolide Chemical compound C1OC(=O)[C@H]2C[C@H](O)[C@H](O)C[C@]2(C)[C@H]2CC[C@]3(C)[C@@H]([C@H](C)[C@H](O)[C@@H](O)[C@H](C)C(C)C)CC[C@H]3[C@@H]21 IXVMHGVQKLDRKH-YEJCTVDLSA-N 0.000 description 1
- 102000005869 Activating Transcription Factors Human genes 0.000 description 1
- 108010005254 Activating Transcription Factors Proteins 0.000 description 1
- 101000910359 Arabidopsis thaliana CDPK-related kinase 1 Proteins 0.000 description 1
- 101100263704 Arabidopsis thaliana VIN3 gene Proteins 0.000 description 1
- 241000894006 Bacteria Species 0.000 description 1
- 241000219198 Brassica Species 0.000 description 1
- 235000003351 Brassica cretica Nutrition 0.000 description 1
- 240000007124 Brassica oleracea Species 0.000 description 1
- 235000011302 Brassica oleracea Nutrition 0.000 description 1
- 235000010149 Brassica rapa subsp chinensis Nutrition 0.000 description 1
- 235000000536 Brassica rapa subsp pekinensis Nutrition 0.000 description 1
- 241000499436 Brassica rapa subsp. pekinensis Species 0.000 description 1
- 235000003343 Brassica rupestris Nutrition 0.000 description 1
- IXVMHGVQKLDRKH-VRESXRICSA-N Brassinolide Natural products O=C1OC[C@@H]2[C@@H]3[C@@](C)([C@H]([C@@H]([C@@H](O)[C@H](O)[C@H](C(C)C)C)C)CC3)CC[C@@H]2[C@]2(C)[C@@H]1C[C@H](O)[C@H](O)C2 IXVMHGVQKLDRKH-VRESXRICSA-N 0.000 description 1
- OYPRJOBELJOOCE-UHFFFAOYSA-N Calcium Chemical compound [Ca] OYPRJOBELJOOCE-UHFFFAOYSA-N 0.000 description 1
- 102000018605 Caldesmon Human genes 0.000 description 1
- 241001249699 Capitata Species 0.000 description 1
- 241000282693 Cercopithecidae Species 0.000 description 1
- 241000033870 Citrullus lanatus subsp. vulgaris Species 0.000 description 1
- 235000012840 Citrullus vulgaris Nutrition 0.000 description 1
- 229920000742 Cotton Polymers 0.000 description 1
- 240000001980 Cucurbita pepo Species 0.000 description 1
- 235000009852 Cucurbita pepo Nutrition 0.000 description 1
- 240000004585 Dactylis glomerata Species 0.000 description 1
- 101100381997 Danio rerio tbc1d32 gene Proteins 0.000 description 1
- 240000003421 Dianthus chinensis Species 0.000 description 1
- LFQSCWFLJHTTHZ-UHFFFAOYSA-N Ethanol Chemical compound CCO LFQSCWFLJHTTHZ-UHFFFAOYSA-N 0.000 description 1
- 241000234642 Festuca Species 0.000 description 1
- 244000044980 Fumaria officinalis Species 0.000 description 1
- 235000006961 Fumaria officinalis Nutrition 0.000 description 1
- 229930191978 Gibberellin Natural products 0.000 description 1
- 241000245654 Gladiolus Species 0.000 description 1
- 244000068988 Glycine max Species 0.000 description 1
- 235000010469 Glycine max Nutrition 0.000 description 1
- 241000219146 Gossypium Species 0.000 description 1
- 108010033040 Histones Proteins 0.000 description 1
- 101000636213 Homo sapiens Transcriptional activator Myb Proteins 0.000 description 1
- 241000209082 Lolium Species 0.000 description 1
- 240000004296 Lolium perenne Species 0.000 description 1
- 241001424413 Lucia Species 0.000 description 1
- 101100381999 Mus musculus Tbc1d32 gene Proteins 0.000 description 1
- 108091028043 Nucleic acid sequence Proteins 0.000 description 1
- 240000007594 Oryza sativa Species 0.000 description 1
- 235000007164 Oryza sativa Nutrition 0.000 description 1
- 240000004371 Panax ginseng Species 0.000 description 1
- 235000005035 Panax pseudoginseng ssp. pseudoginseng Nutrition 0.000 description 1
- 235000003140 Panax quinquefolius Nutrition 0.000 description 1
- 241000220324 Pyrus Species 0.000 description 1
- 240000005498 Setaria italica Species 0.000 description 1
- 241001406921 Squamosa Species 0.000 description 1
- 108010055297 Sterol Esterase Proteins 0.000 description 1
- 101710172711 Structural protein Proteins 0.000 description 1
- 108091023040 Transcription factor Proteins 0.000 description 1
- 102000040945 Transcription factor Human genes 0.000 description 1
- 235000009392 Vitis Nutrition 0.000 description 1
- 241000219095 Vitis Species 0.000 description 1
- 240000008042 Zea mays Species 0.000 description 1
- 235000005824 Zea mays ssp. parviglumis Nutrition 0.000 description 1
- 235000002017 Zea mays subsp mays Nutrition 0.000 description 1
- 230000036579 abiotic stress Effects 0.000 description 1
- 230000009471 action Effects 0.000 description 1
- 238000000246 agarose gel electrophoresis Methods 0.000 description 1
- 230000007152 anther development Effects 0.000 description 1
- 238000013528 artificial neural network Methods 0.000 description 1
- 238000003149 assay kit Methods 0.000 description 1
- 230000009286 beneficial effect Effects 0.000 description 1
- 230000003115 biocidal effect Effects 0.000 description 1
- 230000015572 biosynthetic process Effects 0.000 description 1
- QKSKPIVNLNLAAV-UHFFFAOYSA-N bis(2-chloroethyl) sulfide Chemical compound ClCCSCCCl QKSKPIVNLNLAAV-UHFFFAOYSA-N 0.000 description 1
- 210000000081 body of the sternum Anatomy 0.000 description 1
- 229910052791 calcium Inorganic materials 0.000 description 1
- 239000000969 carrier Substances 0.000 description 1
- 230000036978 cell physiology Effects 0.000 description 1
- 235000013339 cereals Nutrition 0.000 description 1
- 239000003153 chemical reaction reagent Substances 0.000 description 1
- 150000001875 compounds Chemical class 0.000 description 1
- 238000007796 conventional method Methods 0.000 description 1
- 235000005822 corn Nutrition 0.000 description 1
- 238000012258 culturing Methods 0.000 description 1
- 210000004292 cytoskeleton Anatomy 0.000 description 1
- 101150024923 da gene Proteins 0.000 description 1
- 230000006196 deacetylation Effects 0.000 description 1
- 238000003381 deacetylation reaction Methods 0.000 description 1
- 230000004665 defense response Effects 0.000 description 1
- 230000007812 deficiency Effects 0.000 description 1
- 238000000151 deposition Methods 0.000 description 1
- 238000013461 design Methods 0.000 description 1
- 230000008034 disappearance Effects 0.000 description 1
- 239000012153 distilled water Substances 0.000 description 1
- 238000001962 electrophoresis Methods 0.000 description 1
- 210000003527 eukaryotic cell Anatomy 0.000 description 1
- 230000005284 excitation Effects 0.000 description 1
- 239000013613 expression plasmid Substances 0.000 description 1
- 239000004459 forage Substances 0.000 description 1
- 239000012634 fragment Substances 0.000 description 1
- 238000010230 functional analysis Methods 0.000 description 1
- 230000004034 genetic regulation Effects 0.000 description 1
- IXORZMNAPKEEDV-UHFFFAOYSA-N gibberellic acid GA3 Natural products OC(=O)C1C2(C3)CC(=C)C3(O)CCC2C2(C=CC3O)C1C3(C)C(=O)O2 IXORZMNAPKEEDV-UHFFFAOYSA-N 0.000 description 1
- 239000003448 gibberellin Substances 0.000 description 1
- 235000008434 ginseng Nutrition 0.000 description 1
- 238000007654 immersion Methods 0.000 description 1
- 230000010354 integration Effects 0.000 description 1
- 230000003834 intracellular effect Effects 0.000 description 1
- 238000012423 maintenance Methods 0.000 description 1
- 230000004060 metabolic process Effects 0.000 description 1
- 239000004531 microgranule Substances 0.000 description 1
- 230000004048 modification Effects 0.000 description 1
- 238000012986 modification Methods 0.000 description 1
- 230000009456 molecular mechanism Effects 0.000 description 1
- 235000010460 mustard Nutrition 0.000 description 1
- 108091027963 non-coding RNA Proteins 0.000 description 1
- 102000042567 non-coding RNA Human genes 0.000 description 1
- 239000002773 nucleotide Substances 0.000 description 1
- 125000003729 nucleotide group Chemical group 0.000 description 1
- 230000003287 optical effect Effects 0.000 description 1
- 235000002252 panizo Nutrition 0.000 description 1
- 235000021017 pears Nutrition 0.000 description 1
- 230000008288 physiological mechanism Effects 0.000 description 1
- 230000037039 plant physiology Effects 0.000 description 1
- 230000007198 pollen germination Effects 0.000 description 1
- 230000002035 prolonged effect Effects 0.000 description 1
- 230000001737 promoting effect Effects 0.000 description 1
- 230000006916 protein interaction Effects 0.000 description 1
- 238000012797 qualification Methods 0.000 description 1
- 238000011084 recovery Methods 0.000 description 1
- 230000011514 reflex Effects 0.000 description 1
- 230000022532 regulation of transcription, DNA-dependent Effects 0.000 description 1
- 230000001850 reproductive effect Effects 0.000 description 1
- 230000003938 response to stress Effects 0.000 description 1
- 230000011664 signaling Effects 0.000 description 1
- 239000007787 solid Substances 0.000 description 1
- 239000008223 sterile water Substances 0.000 description 1
- 230000001954 sterilising effect Effects 0.000 description 1
- 238000003786 synthesis reaction Methods 0.000 description 1
- 230000002123 temporal effect Effects 0.000 description 1
- 238000012360 testing method Methods 0.000 description 1
- 230000002103 transcriptional effect Effects 0.000 description 1
- 230000009466 transformation Effects 0.000 description 1
- 238000010396 two-hybrid screening Methods 0.000 description 1
- 235000013311 vegetables Nutrition 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
-
- 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]
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)
- Biomedical Technology (AREA)
- Zoology (AREA)
- Biophysics (AREA)
- Biotechnology (AREA)
- General Engineering & Computer Science (AREA)
- Biochemistry (AREA)
- Bioinformatics & Cheminformatics (AREA)
- General Health & Medical Sciences (AREA)
- Wood Science & Technology (AREA)
- Physics & Mathematics (AREA)
- Microbiology (AREA)
- Plant Pathology (AREA)
- Cell Biology (AREA)
- Physiology (AREA)
- Botany (AREA)
- Gastroenterology & Hepatology (AREA)
- Medicinal Chemistry (AREA)
- Proteomics, Peptides & Aminoacids (AREA)
- Breeding Of Plants And Reproduction By Means Of Culturing (AREA)
- Micro-Organisms Or Cultivation Processes Thereof (AREA)
Abstract
本发明公开了一种拟南芥钙调素结合蛋白基因ATIQM2在开花调节中的应用，所述AtIQM2的氨基酸序列经过1个或几个氨基酸残基的取代和/或缺失和或添加且与序列1具有相同功能的有序列1衍生出来的蛋白质。本发明发现ATIQM2基因的编码蛋白能与钙调素结合，其外源DNA插入突变体具有在长日照和短日照条件下延迟开花的表型，说明AtIQM2基因所编码的蛋白具有调节开花的作用。因此，AtIQM2基因可用于控制植物的开花调节，改良植物的经济性状，对农业生产具有重要的意义。
Description
技术领域
本发明涉及基因工程领域，特别是一种拟南芥钙调素结合蛋白基因ATIQM2在开花调节中的应用。
背景技术
高等植物由营养生长向生殖生长转换的过程称为成花诱导，与其生存、繁衍和进化息息相关，其机理尤其是分子机理是当前植物生物学研究的前沿热点。成花诱导过程由遗传和外界环境两个因素决定，受错综复杂的信号途径调控（孙昌辉等，遗传，29：1182，2007）。近年来，在拟南芥中，成花诱导的研究比较系统且进展较快。Srikanth和Schmid（Cell Mol Life Sci，68：2013，2011）在综述中统计了已鉴定的55个基因，加上Braam研究组发现的AtCML23和AtCML24（Delk等，Plant Physiol，139：240，2005；Tsai等，PlantSignal Behav，2：446，2007）共57成花调控基因，分别涉及如下的6个遗传调控途径：光周期途径通过光受体将光信号传给生物钟，再传给主要信号分子CO（CONSTANS），诱导靶基因FT（FLOWERING LOCUS T）的表达，进而实现对成花的调控；CO的时空表达是光周期诱导开花的关键（An等，Development，131：3615，2004）。春化途径的第一阶段是诱导产生的VIN3对FLC（FLOWERING LOCUS C）的去乙酰化（Sung等，Nature，427：159，2004）和非编码RNA对FLC转录的抑制（Swiezewski等，Nature，462：799，2009）；第二阶段通过VRNs对FLC组蛋白H3进行甲基化，降低成花抑制子FLC的水平（De Lucia等，Proc Natl Acad Sci USA，105：16831，2008）。赤霉素途径通过DELLAs和MYBs来调控下游的成花整合基因（Achard等，PlantPhysiol，143：1163，2007）。自主途径的所有基因通过抑制FLC的表达间接地促进成花（Srikanth和Schmid，Cell Mol Life Sci，68：2013，2011）。温度途径主要通过FLC、FLM（FLOWERING LOCUS M）和SVP（SHORT VEGETATIVE PHASE）调节开花（Blázquez等，NatGenet，33：168，2003）。年龄途径独立于其他几条途径，依赖于microRNA156，后者的水平随年龄增加而下降，解除其对靶基因SPL（SQUAMOSA PROMOTER BINDING PROTEIN-LIKE）的抑制而调控开花（Wang等，Cell，138：738，2009）。
在多变的外部环境和内部生理条件下，这57个基因所编码的蛋白通过SOC1、FT和LFY等整合因子实现对开花的精准控制（张素芝和左建儒，2006；Srikanth和Schmid，CellMol Life Sci，68：2013，2011）。尽管如此，有关的研究数据还十分有限，离全面揭示整个成花调控的遗传网络还有很长的路要走。
作为一种重要的第二信使，钙离子（Ca2+）在植物细胞对体内、外刺激所产生的特定生理反应中起着重要作用，参与多种细胞生理和生长发育调控（Yang和Poovaiah，TrendsPlant Sci，8：505，2003）。钙调素（calmodulin，CaM）是包括植物在内的真核细胞内广泛存在的、高度保守的、最重要的一类Ca2+受体，在Ca2+信号转导中起着至关重要的作用。不过，除拟南芥AtCaM7被发现具有转录调控活性（Kushwaha等，Plant Cell，20：1747，2008）外，其他CaM本身不是结构蛋白，也不具备酶、转录因子或离子通道活性，必须通过其下游靶标——CaMBP来调节细胞生理和生长发育。可见，研究CaMBP是解析CaM信号通路的重要途径。然而，已经鉴定和进行过功能分析的植物CaMBP还十分有限，其功能涉及生长发育、代谢调节、细胞骨架功能、离子通道、激素合成与反应、转录调控、胁迫反应等过程（毛国红等，植物生理与分子生物学报，30：481，2004；Bouch é等，Annu Rev Plant Biol，56：435，2005；韦慧彦等，生物化学与生物物理进展，34：124，2007；DeFalco等，Biochem J，425：27，2010；田长恩和周玉萍，植物学报，48：447，2013），例如AtCCaMK参与花药发育（Poovaiah等，Planta，209：161，1999）、AtNPG1参与花粉萌发（Golovkin和Reddy，Proc Natl Acad Sci USA，100：10558，2003）、AtDWF1参与油菜素内酯合成（Du和Poovaih，Nature，437：741，2005）、AtCBK3（Liu等，Plant J，55：760，2008）参与非生物胁迫反应、AtMLO家族（Kim等，Nature，416：447，2002）参与植物防卫反应等。
早在1989年，Friedman等就发现Ca2+参与牵牛的成花调控。不过，在已鉴定的拟南芥成花调控基因中，涉及Ca2+/CaM信号的只有AtCML24及其高度同源的AtCML23。这2个基因编码的钙调素类似蛋白（calmoduolin-like sensor，CAL）均能在与Ca2+结合后发生构象变化；AtCML24表达下调突变体在长日照下延迟10-30天开花（Delk等，Plant Physiol，139：240，2005），而单独AtCML23的功能缺失突变体无成花表型，只有在与同样没有明显成花表型的AtCML24的弱突变体组成双突变体时才表现延迟成花，说明二者在成花调控方面功能冗余且AtCML23的贡献较小；进一步分析发现，这两个蛋白调控包括CO和FLC等成花信号整合基因在内的表达以及调控成花的NO的产生（Tsai等，Plant Signal Behav，2：446，2007）；不过，与它们直接结合的下游靶标仍不清楚。除此以外，仅有烟草的两个钙调素结合蛋白激酶NtMCK1和NtCBK1被发现涉及成花调控（梁述平等，中国科学C辑，31：306，2001；Hua等，JBiol Chem，279：31483，2004）。因此，寻找更多参与成花调控的Ca2+/CaM信号通路成员，研究其分子机理并应用于植物开花时间的控制，具有重要的理论和实践意义。
发明内容
本发明的目的在于提供一种拟南芥钙调素结合蛋白基因ATIQM2。
本发明的另一目的在于提供一种拟南芥钙调素结合蛋白基因ATIQM2在开花调节中的应用。
本发明是通过如下技术方案来实现的：拟南芥钙调素结合蛋白基因ATIQM2在开花调节中的应用，所述拟南芥钙调素结合蛋白基因ATIQM2为以下氨基酸序列组成的蛋白质：Met Gly Val Ser Phe Ser Cys Pro Phe Ala Glu Gln Asp Asp Val Glu Ala Ala LeuAsp Ser Val Thr Val Lys Ser Ile Ser Phe Gly Asp Asp Asp Glu Cys Lys Thr ProLys Arg Ser Val Asn Phe Asn Asp Gly Thr Leu Glu Pro Thr Ile Leu Lys Ser MetGly Ser Gly Lys Met Val Val Glu Lys Ser Val Ser Leu Lys Gly Met Gln Leu GluArg Met Ile Ser Leu Asn Arg Ser Val Lys Asp Asn Gly Phe Glu Ile Ala Lys GluPhe Ser Val Leu Asp Pro Arg Asn Pro Lys His Glu Ala Ala Ile Lys Leu Gln LysVal Tyr Lys Ser Phe Arg Thr Arg Arg Lys Leu Ala Asp Cys Ala Val Leu Val GluGln Ser Trp Trp Lys Leu Leu Asp Phe Ala Glu Leu Lys Arg Ser Ser Ile Ser PhePhe Asp Ile Glu Lys His Glu Thr Ala Ile Ser Arg Trp Ser Arg Ala Arg Thr ArgAla Ala Lys Val Gly Lys Gly Leu Ser Lys Asn Gly Lys Ala Gln Lys Leu Ala LeuGln His Trp Leu Glu Ala Ile Asp Pro Arg His Arg Tyr Gly His Asn Leu His PheTyr Tyr Asn Lys Trp Leu His Cys Gln Ser Arg Glu Pro Phe Phe Tyr Trp Leu AspIle Gly Glu Gly Lys Glu Val Asn Leu Val Glu Lys Cys Pro Arg Leu Lys Leu GlnGln Gln Cys Ile Lys Tyr Leu Gly Pro Met Glu Arg Lys Ala Tyr Glu Val Val ValGlu Asp Gly Lys Phe Phe Tyr Lys His Ser Gly Glu Ile Leu Gln Thr Ser Asp MetGlu Asp Ser Glu Ser Lys Trp Ile Phe Val Leu Ser Thr Ser Lys Val Leu Tyr ValGly Lys Lys Lys Lys Gly Thr Phe Gln His Ser Ser Phe Leu Ala Gly Gly Ala ThrVal Ala Ala Gly Arg Leu Val Val Glu Asn Gly Val Leu Lys Ala Val Trp Pro HisSer Gly His Tyr Gln Pro Thr Glu Glu Asn Phe Met Asp Phe Leu Ser Phe Leu ArgGlu Asn Asp Val Asp Ile Thr Asp Val Lys Met Ser Pro Thr Asp Glu Asp Glu PheSer Ile Tyr Lys Gln Arg Ser Thr His Met Arg Asn His Ser Leu Glu Glu Asp LeuGlu Ala Glu Lys Thr Ile Ser Phe Gln Asp Lys Val Asp Pro Ser Gly Glu Glu GlnThr Leu Met Arg Asn Glu Ser Ile Ser Arg Lys Gln Ser Asp Leu Glu Thr Pro GluLys Met Glu Ser Phe Ser Thr Phe Gly Asp Glu Ile Gln Ser Val Gly Ser Lys SerThr Lys Val Ser Glu Asp Tyr Asp Ser Gly Asp Asp Glu Glu Glu Glu Glu Glu MetPhe Glu Leu Glu Gln Glu Ser Met Pro Ser Glu Gln Ser Ser Pro Arg Gly Glu GluLys Glu Glu Gly Glu Thr Lys Glu Ser Glu Val Val Lys Ile Thr Glu Glu Ser IleLeu Lys Arg Ile Asn Ser Lys Lys Glu Thr Lys Ser Phe Gln Leu Gly Lys Gln LeuSer Cys Lys Trp Thr Thr Gly Ala Gly Pro Arg Ile Gly Cys Val Arg Asp Tyr ProSer Glu Leu Gln Phe Gln Ala Leu Glu Gln Val Asn Leu Ser Pro Arg Ser Ala SerVal Ser Arg Leu Cys Phe Ser Ser Ser Ser Gln Thr Gln Thr Pro Gln Met Ser ProLeu Trp Arg Gly Met Ser Leu Pro Thr Asp Ile Thr Leu Thr Asn Ser。
上述氨基酸序列经过1个或几个氨基酸残基的取代和/或缺失和或添加且与上述氨基酸序列具有相同功能的上述氨基酸序列衍生的蛋白质。
所述拟南芥钙调素结合蛋白基因ATIQM2的基因序列为：
atgggtgtatcattctcctgtccgttcgcagagcaagacgacgtggaagctgctctagactctgtcacggttaagtccataagctttggcgatgacgacgaatgcaaaactcctaagagatctgttaatttcaatgacggtactttagagcccacgattttgaaatctatgggatctggtaaaatggttgttgagaaatctgttagcttgaaagggatgcagctagagagaatgatctcacttaacaggtctgttaaagacaatggctttgagattgctaaagagttctcagttcttgatccgaggaatccgaaacacgaagctgctattaagttgcagaaggtttataagagctttcgtactagacgaaagctcgctgattgcgctgtgcttgttgagcagagctggtggaaacttttggattttgctgagttgaagagaagttctatatctttctttgatattgagaaacacgaaaccgcgatctcaaggtggtctagagcaagaactagagcagctaaagttggtaaaggtttgtcaaagaatggaaaagctcaaaagcttgctttacaacactggcttgaagcgattgacccgagacatcggtatggacacaacctgcacttttactacaacaaatggctccattgtcagagtagagaacctttcttctactggcttgatatcggcgaggggaaagaggtaaatcttgtggagaaatgtccgcggttgaaacttcaacaacagtgtatcaaataccttggtccgatggaaagaaaagcttatgaggtggttgtggaagacggcaaattcttctacaagcatagtggagagattcttcaaacttctgatatggaagatagtgaatccaaatggatttttgtgctaagcacatcgaaagtgttatacgtcgggaagaagaagaagggtacgtttcaacattcaagcttcttagctggaggagctactgttgctgcagggagattagttgttgagaatggtgttcttaaggctgtttgg ccacatagcggacattatcaacctacagaagagaatttcatggactttctctctttcctccgcgagaatgatgtcgatatcaccgatgtaaagatgagtcctacagatgaagatgaattctctatttacaaacagagaagcactcatatgagaaaccattctttagaagaggatttggaggctgagaagaccatttcctttcaagataaggttgatccaagtggagaagaacaaactctgatgagaaatgagtccatctcaagaaagcaaagcgatctcgagacaccggaaaagatggaatcatttagcacatttggcgatgaaatccaatccgtgggatcgaagtctacaaaagtatctgaagactacgattcaggtgacgatgaagaggaagaagaagagatgtttgagttagaacaagaatcaatgccttcagagcaaagctcacctagaggagaagaaaaagaagaaggagaaaccaaggagagtgaagtagtgaagataacagaagaatcaatcctaaaaaggatcaattcaaagaaggaaactaaatctttccaacttgggaaacaactatcatgcaaatggacaacaggtgcaggaccaagaatcggttgtgtaagagattatccatcagagcttcagtttcaagcactggaacaagtgaacttgtctccgagaagtgcttctgtttcaagactctgtttctcttcttcatcgcaaacgcaaacgcctcaaatgtcaccgttatggcgaggaatgtcactacctacagatatcacactcacgaattcataa
限定的DNA序列具有90%以上的同源性，且编码相同功能蛋白的DNA分子。
所述基因的编码序列的DNA分子来控制植物的开花时间。
通过使用所述控制植物的开花时间的基因来控制植物的开花时间。
开花时间通过在植物中过度表达拟南芥钙调素结合蛋白基因ATIQM2分子而被加速。
开花时间通过抑制拟南芥钙调素结合蛋白基因ATIQM2的DNA分子的表达而被延迟。
所述植物包括水稻、小麦、大麦、玉米、大豆、马铃薯、红豆、燕麦和粟等在内的粮食作物；包括拟南芥、大白菜、萝卜、辣椒、草莓、番茄、西瓜、黄瓜、卷心菜、甜瓜、西葫芦、韭菜、洋葱和胡萝卜等在内的蔬菜作物；包括人参、烟草、棉花、芝麻、甘蔗、甜菜、野生芝麻、花生和油菜籽等在内的经济作物；包括苹果、梨、枣、桃、称猴桃、葡萄、桔子、柿子、李子、杏和香蕉等在内的水果；包括玫瑰、剑兰、大丁草、康乃馨、菊花、百合和郁金香等 在内的花卉；和包括黑麦草、红三叶草、鸭茅、紫花首楷、高酥油草和多年生黑麦草等在内的饲料作物。
在研究拟南芥中与控制开花时间的新基因时，本发明人发现了AtIQM2所编码蛋白AtIQM2在酵母和洋葱表皮细胞中能与钙调素ATCaM5结合蛋白（见图1），其突变体——iqm2-1和iqm2-2（见图2）均呈现出长日照（16h光照+8h黑暗）和短日照条件下（8h光照+16h黑暗）延迟开花的表型（见图3），说明ATIQM2是一个钙调素结合蛋白，具有促进植物开花的功能。因此，可通过基因工程手段，促进或抑制AtIQM2基因的表达，进而调节植物的开花时间。
与现有技术相比，本发明的有益效果：既可通过加速植物开花在短时间内产生花和种子，还可通过延迟植物开花维持营养生长，改善植物有用部位（如叶片或茎）的生产效率。因此，本发明对农业生产具有重要意义。
附图说明
图1为拟南芥ATIQM2基因的编码蛋白ATIQM2的钙调素结合分析结果图（其中，A）示ATIQM2蛋白与ATCaM5的酵母双杂交分析结果【左为完全培养基；右为营养缺陷培养基（有菌为阳性结果）】：1为pHybLex/Zeo空载体，阴性对照；1A为pHybLex/Zeo+pYESTrp2，阴性对照；1B为pHybLex/Zeo+pYESTrp2-IQM2，阴性对照；2A为pHybLex/Zeo-CaM5+pYESTrp2，阴性对照；2B为pHybLex/Zeo-CaM5+pYESTrp2-IQM2，样品检测；3C为pHybLex/Zeo-Fos2+pYESTrp2-Jun，阳性对照。B）示ATIQM2蛋白与ATCaM5的双分子荧光互补（BiFC）分析结果。上行图为荧光显微镜照片；下行图为与荧光观察相同材料的相差显微镜照片，箭头所指为细胞核。左（IQM2）为pSATN-nEYFP-IQM2+pSATN-cEYFP，阴性对照；中（CaM5）为pSATN-nEYFP+pSATN-cEYFP-CaM2，阴性对照；右（IQM2+CaM5）为pSATN-nEYFP-IQM2+pSATN-cEYFP-CaM2）。
图2为拟南芥ATIQM2基因的插入突变体的鉴定结果图（其中，A）示拟南芥ATIQM2的结构及其突变体iqm2-1和iqm2-2的T-DNA和转座子的插入位点，iqm2-1的遗传背景为Columbia生态型；iqm2-2的遗传背景为Landsberg生态型。B）示ATIQM2基因的插入突变体的ATIQM2基因表达的RT-PCR检测结果。 途中1、2、3、4分别代表iqm2-1、iqm2-2、Columbia和Landsberg，M为DNA分子量标准。左图为管家基因β-ATPase的RT-PCR分析结果，电泳条带基本一致，说明所用RNA量一致；右图为突变体iqm2-1和iqm2-2及其相应的野生型的RT-PCR分析结果，在两个野生型能检测出而在两个突变体中未能检测出ATIQM2基因的表达产品——mRNA，说明iqm2-1和iqm2-2均为功能缺失突变体）。
图3为拟南芥野生型与ATIQM2基因插入突变体在长日照（每天光照16小时，黑暗8小时）条件下开花表型的观察结果图（其中，A）示代表性植物的照片。B）示突变体iqm2-2及其野生型植株Landsberg的开花的时间进程。C）示开花时突变体iqm2-2及其野生型植株Landsberg的莲座叶数，叶片多表示迟开花）。
具体实施方式
下面结合附图和实施例对本发明做进一步说明，但不是以任何形式限制本发明。
下面结合具体实施例对本发明做进一步详细的描述，这些实施例只用来说明本发明，并不限制本发明的范围。
实施例中所用的方法如无特别的说明，均为常规方法；所用材料如无特别的说明，均购自常规生化试剂公司；引物合成由宝生物工程（大连）有限公司完成，测序工作由华大基因科技服务有限公司完成。
本发明所提供的ATIQM2基因的核酸序列如SEQ ID No:2（序列2）。ATIQM2基因编码的蛋白质，命名为ATIQM2蛋白质，其氨基酸序列如SEQ ID No:1（序列1）。
实施例1拟南芥ATIQM2的钙调素结合活性分析
生物信息学分结果显示，ATIQM2蛋白的N端含有1个IQ基序，因此可能是一个钙调素结合蛋白。为了探究ATIQM1蛋白是否具有钙调素结合特性，特进行如下所述的酵母双杂交实验和双分子荧光互补（BiFC）实验。
一、酵母双杂交实验
酵母双杂交实验是一种直观、快速地判断两个目标蛋白在酵母细胞中相互 作用的新技术，将待研究的两种蛋白质的基因分别克隆到酵母表达质粒的转录激活因子（如GAL4等）的DNA结合结构域编码序列和GAL4激活结构域编码序列，构建成融合表达载体，从表达产物来分析两种蛋白质相互作用。
首先，构建用于酵母双杂交实验的表达载体。
其一，利用ATIQM2基因的带限制性酶切位点的特异引物ATIQM2-F1和ATIQM2-R1，对从拟南芥幼苗提取的总RNA进行RT-PCR扩增，得到两端分别具有BamH I和Not I的ATIQM2基因的CDS，外送测序公司；然后，将经过测序证实的ATIQM2基因的CDS通过BamH I/Not I位点定向克隆入用于酵母双杂交的捕获物（prey）载体pYESTrp2中。构建捕获物（prey）载体pYESTrp2所用ATIQM2基因的带限制性酶切位点的特异引物如下：
ATIQM2-F1:5’-GGATCCATGGGTGTATCATTCTCCT3-3’
BamH I
ATIQM2-R1:5’-GCGGCCGCACTTCTCTTCAACTCAGC-3’
Not I
其二，利用ATCaM5基因的带限制性酶切位点的特异引物ATCaM5-F1和ATCaM5-R1，对从拟南芥幼苗提取的总RNA进行RT-PCR扩增，得到两端分别具有EcoR I和Not I的ATCaM5基因的CDS，外送测序公司；然后，将经过测序证实的ATCaM5基因的CDS通过EcoR I/Not I位点定向克隆入用于酵母双杂交的诱饵（bait）载体pHybLex/Zeo中。构建诱饵（bait）载体pHybLex/Zeo-CaM5所用的ATCaM5基因的带限制性酶切位点的特异引物如下：
ATCaM5-F1:5’-CGCGAATTCATGGCAGATCAGCTCACCGATGATC-3’
EcoR I
ATCaM5-R1:5’-AGAGCGGCCGCTCACTTTGCCATCATAACTTTGAC-3’
Not I
其次，进行酵母双杂交实验。
在完成上述两种载体的构建后，以pHybLex/Zeo-CaM5为诱饵，以pYESTrp2-IQM2为捕获物，将不同组合质粒分两步共转化感受态的酵母细胞L40， 经营养缺陷型筛选和抗生素的抗性筛选，得到含有两个转化质粒的酵母细胞，同时设置必要的阴性对照和阳性对照，进行ATIQM2蛋白和ATCaM5蛋白的酵母双杂交实验，结果如图1A所示。
图2A左为含不同质粒组合的酵母细胞在完全培养基（YPAD）平板上的生长情况，结果显示各种组合均能正常生长，没有差异；图1A右为含不同质粒组合的酵母细胞在营养缺陷和显色平板（YC-WKUK）平板上的生长情况，结果显示各种阴性对照组合的酵母细胞不能生长。上述酵母双杂交实验的结果表明，ATIQM2在酵母中能与钙调素ATCaM5结合。
二、双分子荧光互补（BiFC）实验
尽管酵母双杂交实验结果显示ATIQM2在酵母中能与钙调素ATCaM5结合，但在植物细胞中ATIQM2未必能就与钙调素ATCaM5的结合。因此，进一步开展双分子荧光互补（BiFC）实验加以验证。
BiFC是一种直观、快速地判断两个目标蛋白在活细胞中相互作用的新技术，是指两个不发光的荧光蛋白互补片段在与其融合蛋白质的相互作用的驱动下重新组装形成荧光复合物，恢复荧光特性。如果荧光蛋白活性恢复则表明两目标蛋白发生了相互作用。
首先，构建用于BiFC实验的表达载体。
其一，利用ATIQM2基因的带限制性酶切位点的特异引物ATIQM2-F2和ATIQM2-R2，对从拟南芥幼苗提取的总RNA进行RT-PCR扩增，得到两端分别具有Sal I和BamH I的ATIQM2基因的CDS，外送测序公司；然后，将经过测序证实的ATIQM2基因的CDS通过Sal I/BamH I位点定向克隆入用于BiFC的EYFP的N-端载体pSATN-nEYFP的nEYFP下游，生产融合蛋白nEYFP-IQM2。构建载体pSATN-nEYFP-IQM2所用ATIQM2基因的带限制性酶切位点的特异引物如下：
ATIQM2-F2:5’-GTCGACATGGGTGTATCATTCTCCT3-3’
SalI
ATIQM2-R2:5’-GGATCCACTTCTCTTCAACTCAGC-3’
BamH I
其二，利用ATCaM5基因的带限制性酶切位点的特异引物ATCaM5-F2和ATCaM5-R2，对从拟南芥幼苗提取的总RNA进行RT-PCR扩增，得到两端分别具有EcoR I和Sal I的ATCaM5基因的CDS，外送测序公司测序；然后，将经过测序证实的ATCaM5基因的CDS通过EcoR I/SalI位点定向克隆入用于BiFC的EYFP的C-端载体pSATN-cEYFP的cEYFP下游，生产融合蛋白cEYFP-CaM5。构建载体pSATN-cEYFP-CaM5所用ATIQM2基因的带限制性酶切位点的特异引物如下：
ATCaM5-F2:5’-CGCGAATTCATGGCAGATCAGCTCACCGATGATC-3’
EcoR I
ATCaM5-R2:5’-AGAGTCGACTCACTTTGCCATCATAACTTTGAC-3’
Sal I
其次，进行BiFC实验。
其一，大量制备2种对照空质粒pSATN-nEYFP和pSATN-cEYFP以及和2种构建的基因融合质粒pSATN-nEYFP-IQM2和pSATN-cEYFP-CaM2。其二，根据Helios Gene Gun System提供的方法制备用于基因枪转化的带有上述各种质粒的微粒子弹；其三，用基因枪将上述子弹，轰击进平铺在MS培养基平板表面黑暗下25℃培养16小时的洋葱鳞茎内表皮细胞内，继续在25℃黑暗培养；其四，培养16小时后，利用荧光显微镜对洋葱鳞茎内表皮细胞进行观察，实验结果显示，在蓝色激发光作用下，同一张表皮细胞中有多个洋葱表皮细胞产生出较强的荧光，而且荧光主要分布在细胞核（图1下右）；但对照只有均匀、非常微弱的自发荧光（图1下左、中）。可见，ATIQM2和ATCaM2能够在植物细胞内结合。
上述酵母双杂交和双分子荧光互补实验的结果都说明ATIQM2是1个钙调素结合蛋白。
实施例2拟南芥ATIQM2突变体的鉴定
T-DNA插入突变体iqm2-1（SALK-137935.25.90.x）种子购自ABRC（theArabidopsis Biological Resource Center,Ohio State University,USA）。 转座子插入突变体iqm2-2（GT21916）种子由CSHL（Cold Spring Harbor Laboratory,New York onLong Island,USA）提供。ABRC提供的Salk T-DNA插入资料显示，iqm2-1的T-DNA反向插在ATIQM2的第1个外显子的+370bp处；TAIR（the arabidopsis information resource）iqm2-2的转座子DS5-4正向插入第1个外显子的+26bp处中（图2A）。据此，我们对相应片段进行了测序，结果证实确实如此。
一般而言，从种子库中购买的种子有可能含有杂合子及野生型的植株，所以需要对所购种子进行分子标记鉴定，从中筛选出纯合子。
将突变体iqm2-1和iqm2-2和相应野生型Columbia和Landsberg的种子消毒灭菌后分别接种在1/2MS固体培养基上，光下培养10天，随机取12株幼苗，利用Trizol试剂盒提取植物总RNA，然后用One-Step RT-PCR试剂盒进行扩增，所用引物如下：
ATIQM2引物：其在基因的相应位置见图2A，预期产物长度为1815bp
正向引物:ATGGGTGTATCATTCTCCTGTC
反向引物:TTATGAATTCGTGAGTGTGATATC
β-ATPase：预期产物长度为333bp
正向引物:CTGAATCAATCTCTTAAGCTGCTG
反向引物:GTGCAGAAAGTTCTACAGAACTAC
将RT-PCR扩增产物进行琼脂糖凝胶电泳，结果如图2B所示。图2B左为突变体iqm2-1和iqm2-2及其相应的野生型中看家基因β-ATPase的RT-PCR分析结果，电泳条带大小与预期的333bp一致，每个条带的宽度和亮度基本一致，说明所用RNA量基本一致；图2B右为突变体iqm2-1和iqm2-2及其相应的野生型中ATIQM2基因的RT-PCR分析结果，在两个野生型（3和4）中能检测出ATIQM2基因的表达产品——mRNA，而在两个突变体（1和2）中未能检出所预期产品，说明iqm2-1和iqm2-2的ATIQM2基因因为外源DNA的插入而被破坏，成为功能缺失突变体，可以作为ATIQM2基因功能研究的材料。
实施例3拟南芥AtIQM2参与开花调控的发现
功能缺失突变体是进行基因功能分析的良好材料。本发明发明人鉴定获得了两个不同生态型的AtIQM2的功能缺失突变体iqm2-1和iqm2-2。为了进一步了解ATIQM2基因在植株生长发育过程中的作用，拟通过实验观察了突变体iqm2-1及其野生型Columbia以及iqm2-2及其野生型Landsberg的生长情况。
将前述4种拟南芥种子用适量蒸馏水浸泡后，在4℃冰箱处理48小时，然后用70%乙醇浸泡30s，再用0.1%HgCl2消毒灭菌5分钟，最后用无菌水漂洗3次，播种至经过高温灭菌的营养土与蛭石比例为1︰1的土壤中，置于16小时光照/8小时黑暗的光周期、800mW/m2的光照强度、70-80%的相对湿度、22-24℃的温度条件下培养。每隔3天浇水一次，6天施肥一次。逐日观察各基因型植株的生长发育情况，最典型的差异是突变体与野生型的开花时间不同（图3A）。以本领域熟知的拟南芥花茎长度达1厘米时为开花标志，与野生型植株Columbia和Landsberg相比，突变体iqm2-1和iqm2-2晚开花2-3天，开花时植株的莲座叶数分别多1-2片或2-3片（图3）。因为两个不同生态型拟南芥的ATIQM2基因的功能缺失突变体iqm2-1和iqm2-2都显示出开花延迟的表型，说明ATIQM2基因所编码的蛋白ATIQM2具有促进开花的功能。
以上详细描述了本发明的较佳具体实施例，应当理解，本领域的普通技术无需创造性劳动就可以根据本发明的构思做出诸多修改和变化。因此，凡本技术领域中技术人员依本发明构思在现有技术基础上通过逻辑分析、推理或者根据有限的实验可以得到的技术方案，均应该在由本权利要求书所确定的保护范围之中。
<110> 广州大学
<120> 拟南芥钙调素结合蛋白基因ATIQM2在开花调节中的应用
<160> 14
<210> 1
<211> 1815
<212> RNA
<213> 未知
<220>
<223>
<400>1
Met Gly Val Ser Phe Ser Cys Pro Phe Ala Glu Gln Asp Asp Val Glu AlaAla Leu Asp Ser Val Thr Val Lys Ser Ile Ser Phe Gly Asp Asp Asp Glu Cys LysThr Pro Lys Arg Ser Val Asn Phe Asn Asp Gly Thr Leu Glu Pro Thr Ile Leu LysSer Met Gly Ser Gly Lys Met Val Val Glu Lys Ser Val Ser Leu Lys Gly Met GlnLeu Glu Arg Met Ile Ser Leu Asn Arg Ser Val Lys Asp Asn Gly Phe Glu Ile AlaLys Glu Phe Ser Val Leu Asp Pro Arg Asn Pro Lys His Glu Ala Ala Ile Lys LeuGln Lys Val Tyr Lys Ser Phe Arg Thr Arg Arg Lys Leu Ala Asp Cys Ala Val LeuVal Glu Gln Ser Trp Trp Lys Leu Leu Asp Phe Ala Glu Leu Lys Arg Ser Ser IleSer Phe Phe Asp Ile Glu Lys His Glu Thr Ala Ile Ser Arg Trp Ser Arg Ala ArgThr Arg Ala Ala Lys Val Gly Lys Gly Leu Ser Lys Asn Gly Lys Ala Gln Lys LeuAla Leu Gln His Trp Leu Glu Ala Ile Asp Pro Arg His Arg Tyr Gly His Asn LeuHis Phe Tyr Tyr Asn Lys Trp Leu His Cys Gln Ser Arg Glu Pro Phe Phe Tyr TrpLeu Asp Ile Gly Glu Gly Lys Glu Val Asn Leu Val Glu Lys Cys Pro Arg Leu LysLeu Gln Gln Gln Cys Ile Lys Tyr Leu Gly Pro Met Glu Arg Lys Ala Tyr Glu ValVal Val Glu Asp Gly Lys Phe Phe Tyr Lys His Ser Gly Glu Ile Leu Gln Thr SerAsp Met Glu Asp Ser Glu Ser Lys Trp Ile Phe Val Leu Ser Thr Ser Lys Val LeuTyr Val Gly Lys Lys Lys Lys Gly Thr Phe Gln His Ser Ser Phe Leu Ala Gly GlyAla Thr Val Ala Ala Gly Arg Leu Val Val Glu Asn Gly Val Leu Lys Ala Val TrpPro His Ser Gly His Tyr Gln Pro Thr Glu Glu Asn Phe Met Asp Phe Leu Ser PheLeu Arg Glu Asn Asp Val Asp Ile Thr Asp Val Lys Met Ser Pro Thr Asp Glu AspGlu Phe Ser Ile Tyr Lys Gln Arg Ser Thr His Met Arg Asn His Ser Leu Glu GluAsp Leu Glu Ala Glu Lys Thr Ile Ser Phe Gln Asp Lys Val Asp Pro Ser Gly GluGlu Gln Thr Leu Met Arg Asn Glu Ser Ile Ser Arg Lys Gln Ser Asp Leu Glu ThrPro Glu Lys Met Glu Ser Phe Ser Thr Phe Gly Asp Glu Ile Gln Ser Val Gly SerLys Ser Thr Lys Val Ser Glu Asp Tyr Asp Ser Gly Asp Asp Glu Glu Glu Glu GluGlu Met Phe Glu Leu Glu Gln Glu Ser Met Pro Ser Glu Gln Ser Ser Pro Arg GlyGlu Glu Lys Glu Glu Gly Glu Thr Lys Glu Ser Glu Val Val Lys Ile Thr Glu GluSer Ile Leu Lys Arg Ile Asn Ser Lys Lys Glu Thr Lys Ser Phe Gln Leu Gly LysGln Leu Ser Cys Lys Trp Thr Thr Gly Ala Gly Pro Arg Ile Gly Cys Val Arg AspTyr Pro Ser Glu Leu Gln Phe Gln Ala Leu Glu Gln Val Asn Leu Ser Pro Arg SerAla Ser Val Ser Arg Leu Cys Phe Ser Ser Ser Ser Gln Thr Gln Thr Pro Gln MetSer Pro Leu Trp Arg Gly Met Ser Leu Pro Thr Asp Ile Thr Leu Thr Asn Ser
<210> 2
<211> 1818
<212> DNA
<213> 未知
<220>
<223>
<400>2
atgggtgtatcattctcctgtccgttcgcagagcaagacgacgtggaagctgctctagactctgtcacggttaagtccataagctttggcgatgacgacgaatgcaaaactcctaagagatctgttaatttcaatgacggtactttagagcccacgattttgaaatctatgggatctggtaaaatggttgttgagaaatctgttagcttgaaagggatgcagctagagagaatgatctcacttaacaggtctgttaaagacaatggctttgagattgctaaagagttctcagttcttgatccgaggaatccgaaacacgaagctgctattaagttgcagaaggtttataagagctttcgtactagacgaaagctcgctgattgcgctgtgcttgttgagcagagctggtggaaacttttggattttgctgagttgaagagaagttctatatctttctttgatattgagaaacacgaaaccgcgatctcaaggtggtctagagcaagaactagagcagctaaagttggtaaaggtttgtcaaagaatggaaaagctcaaaagcttgctttacaacactggcttgaagcgattgacccgagacatcggtatggacacaacctgcacttttactacaacaaatggctccattgtcagagtagagaacctttcttctactggcttgatatcggcgaggggaaagaggtaaatcttgtggagaaatgtccgcggttgaaacttcaacaacagtgtatcaaataccttggtccgatggaaagaaaagcttatgaggtggttgtggaagacggcaaattcttctacaagcatagtggagagattcttcaaacttctgatatggaagatagtgaatccaaatggatttttgtgctaagcacatcgaaagtgttatacgtcgggaagaagaagaagggtacgtttcaacattcaagcttcttagctggaggagctactgttgctgcagggagattagttgttgagaatggtgttcttaaggctgtttggccacatagcggacattatcaacctacagaagagaatttcatggactttctctctttcctccgcgagaatgatgtcgatatcaccgatgtaaagatgagtcctacagatgaagatgaattctctatttacaaacagagaagcactcatatgagaaaccattctttagaagaggatttggaggctgagaagaccatttcctttcaagataaggttgatccaagtggagaagaacaaactctgatgagaaatgagtccatctcaagaaagcaaagcgatctcgagacaccggaaaagatggaatcatttagcacatttggcgatgaaatccaatccgtgggatcgaagtctacaaaagtatctgaagactacgattcaggtgacgatgaagaggaagaagaagagatgtttgagttagaacaagaatcaatgccttcagagcaaagctcacctagaggagaagaaaaagaagaaggagaaaccaaggagagtgaagtagtgaagataacagaagaatcaatcctaaaaaggatcaattcaaagaaggaaactaaatctttccaacttgggaaacaactatcatgcaaatggacaacaggtgcaggaccaagaatcggttgtgtaagagattatccatcagagcttcagtttcaagcactggaacaagtgaacttgtctccgagaagtgcttctgtttcaagactctgtttctcttcttcatcgcaaacgcaaacgcctcaaatgtcaccgttatggcgaggaatgtcactacctacagatatcacactcacgaattcataa
<210> 3
<211> 26
<212> DNA
<213> 人工序列
<220>
<223>
<400>3
5’-GGATCCATGGGTGTATCATTCTCCT-3’
<210> 4
<211> 26
<212> DNA
<213> 人工序列
<220>
<223>
<400>4
5’-GCGGCCGCACTTCTCTTCAACTCAGC-3’
<210> 5
<211> 34
<212> DNA
<213> 人工序列
<220>
<223>
<400>5
5’-CGCGAATTCATGGCAGATCAGCTCACCGATGATC-3’
<210> 6
<211> 35
<212> DNA
<213> 人工序列
<220>
<223>
<400>6
5’-AGAGCGGCCGCTCACTTTGCCATCATAACTTTGAC-3’
<210> 7
<211> 25
<212> DNA
<213> 人工序列
<220>
<223>
<400>7
5’-GTCGACATGGGTGTATCATTCTCCT-3’
<210> 8
<211> 24
<212> DNA
<213> 人工序列
<220>
<223>
<400>8
5’-GGATCCACTTCTCTTCAACTCAGC-3’
<210> 9
<211> 34
<212> DNA
<213> 人工序列
<220>
<223>
<400>9
5’-CGCGAATTCATGGCAGATCAGCTCACCGATGATC-3’
<210>10
<211>33
<212> DNA
<213> 人工序列
<220>
<223>
<400>10
5’-AGAGTCGACTCACTTTGCCATCATAACTTTGAC-3’
<210>11
<211>22
<212> DNA
<213> 人工序列
<220>
<223>
<400>11
ATGGGTGTATCATTCTCCTGTC
<210>12
<211>24
<212> DNA
<213> 人工序列
<220>
<223>
<400>12
TTATGAATTCGTGAGTGTGATATC
<210>13
<211>24
<212> DNA
<213> 人工序列
<220>
<223>
<400>13
CTGAATCAATCTCTTAAGCTGCTG
<210>14
<211>24
<212> DNA
<213> 人工序列
<220>
<223>
<400>14
GTGCAGAAAGTTCTACAGAACTAC。
Claims (6)
1.拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，所述拟南芥钙调素结合蛋白基因ATIQM2所表达的蛋白质由以下氨基酸序列组成：Met Gly Val Ser Phe SerCys Pro Phe Ala Glu Gln Asp Asp Val Glu Ala Ala Leu Asp Ser Val Thr Val LysSer Ile Ser Phe Gly Asp Asp Asp Glu Cys Lys Thr Pro Lys Arg Ser Val Asn PheAsn Asp Gly Thr Leu Glu Pro Thr Ile Leu Lys Ser Met Gly Ser Gly Lys Met ValVal Glu Lys Ser Val Ser Leu Lys Gly Met Gln Leu Glu Arg Met Ile Ser Leu AsnArg Ser Val Lys Asp Asn Gly Phe Glu Ile Ala Lys Glu Phe Ser Val Leu Asp ProArg Asn Pro Lys His Glu Ala Ala Ile Lys Leu Gln Lys Val Tyr Lys Ser Phe ArgThr Arg Arg Lys Leu Ala Asp Cys Ala Val Leu Val Glu Gln Ser Trp Trp Lys LeuLeu Asp Phe Ala Glu Leu Lys Arg Ser Ser Ile Ser Phe Phe Asp Ile Glu Lys HisGlu Thr Ala Ile Ser Arg Trp Ser Arg Ala Arg Thr Arg Ala Ala Lys Val Gly LysGly Leu Ser Lys Asn Gly Lys Ala Gln Lys Leu Ala Leu Gln His Trp Leu Glu AlaIle Asp Pro Arg His Arg Tyr Gly His Asn Leu His Phe Tyr Tyr Asn Lys Trp LeuHis Cys Gln Ser Arg Glu Pro Phe Phe Tyr Trp Leu Asp Ile Gly Glu Gly Lys GluVal Asn Leu Val Glu Lys Cys Pro Arg Leu Lys Leu Gln Gln Gln Cys Ile Lys TyrLeu Gly Pro Met Glu Arg Lys Ala Tyr Glu Val Val Val Glu Asp Gly Lys Phe PheTyr Lys His Ser Gly Glu Ile Leu Gln Thr Ser Asp Met Glu Asp Ser Glu Ser LysTrp Ile Phe Val Leu Ser Thr Ser Lys Val Leu Tyr Val Gly Lys Lys Lys Lys GlyThr Phe Gln His Ser Ser Phe Leu Ala Gly Gly Ala Thr Val Ala Ala Gly Arg LeuVal Val Glu Asn Gly Val Leu Lys Ala Val Trp Pro His Ser Gly His Tyr Gln ProThr Glu Glu Asn Phe Met Asp Phe Leu Ser Phe Leu Arg Glu Asn Asp Val Asp IleThr Asp Val Lys Met Ser Pro Thr Asp Glu Asp Glu Phe Ser Ile Tyr Lys Gln ArgSer Thr His Met Arg Asn His Ser Leu Glu Glu Asp Leu Glu Ala Glu Lys Thr IleSer Phe Gln Asp Lys Val Asp Pro Ser Gly Glu Glu Gln Thr Leu Met Arg Asn GluSer Ile Ser Arg Lys Gln Ser Asp Leu Glu Thr Pro Glu Lys Met Glu Ser Phe SerThr Phe Gly Asp Glu Ile Gln Ser Val Gly Ser Lys Ser Thr Lys Val Ser Glu AspTyr Asp Ser Gly Asp Asp Glu Glu Glu Glu Glu Glu Met Phe Glu Leu Glu Gln GluSer Met Pro Ser Glu Gln Ser Ser Pro Arg Gly Glu Glu Lys Glu Glu Gly Glu ThrLys Glu Ser Glu Val Val Lys Ile Thr Glu Glu Ser Ile Leu Lys Arg Ile Asn SerLys Lys Glu Thr Lys Ser Phe Gln Leu Gly Lys Gln Leu Ser Cys Lys Trp Thr ThrGly Ala Gly Pro Arg Ile Gly Cys Val Arg Asp Tyr Pro Ser Glu Leu Gln Phe GlnAla Leu Glu Gln Val Asn Leu Ser Pro Arg Ser Ala Ser Val Ser Arg Leu Cys PheSer Ser Ser Ser Gln Thr Gln Thr Pro Gln Met Ser Pro Leu Trp Arg Gly Met SerLeu Pro Thr Asp Ile Thr Leu Thr Asn Ser。
2.如权利要求1所述的拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，其特征在于，所述拟南芥钙调素结合蛋白基因ATIQM2的基因序列为：
Atgggtgtatcattctcctgtccgttcgcagagcaagacgacgtggaagctgctctagactctgtcacggttaagtccataagctttggcgatgacgacgaatgcaaaactcctaagagatctgttaatttcaatgacggtactttagagcccacgattttgaaatctatgggatctggtaaaatggttgttgagaaatctgttagcttgaaagggatgcagctagagagaatgatctcacttaacaggtctgttaaagacaatggctttgagattgctaaagagttctcagttcttgatccgaggaatccgaaacacgaagctgctattaagttgcagaaggtttataagagctttcgtactagacgaaagctcgctgattgcgctgtgcttgttgagcagagctggtggaaacttttggattttgctgagttgaagagaagttctatatctttctttgatattgagaaacacgaaaccgcgatctcaaggtggtctagagcaagaactagagcagctaaagttggtaaaggtttgtcaaagaatggaaaagctcaaaagcttgctttacaacactggcttgaagcgattgacccgagacatcggtatggacacaacctgcacttttactacaacaaatggctccattgtcagagtagagaacctttcttctactggcttgatatcggcgaggggaaagaggtaaatcttgtggagaaatgtccgcggttgaaacttcaacaacagtgtatcaaataccttggtccgatggaaagaaaagcttatgaggtggttgtggaagacggcaaattcttctacaagcatagtggagagattcttcaaacttctgatatggaagatagtgaatccaaatggatttttgtgctaagcacatcgaaagtgttatacgtcgggaagaagaagaagggtacgtttcaacattcaagcttcttagctggaggagctactgttgctgcagggagattagttgttgagaatggtgttcttaaggctgtttggccacatagcggacattatcaacctacagaagagaatttcatggactttctctctttcctccgcgagaatgatgtcgatatcaccgatgtaaagatgagtcctacagatgaagatgaattctctatttacaaacagagaagcactcatatgagaaaccattctttagaagaggatttggaggctgagaagaccatttcctttcaagataaggttgatccaagtggagaagaacaaactctgatgagaaatgagtccatctcaagaaagcaaagcgatctcgagacaccggaaaagatggaatcatttagcacatttggcgatgaaatccaatccgtgggatcgaagtctacaaaagtatctgaagactacgattcaggtgacgatgaagaggaagaagaagagatgtttgagttagaacaagaatcaatgccttcagagcaaagctcacctagaggagaagaaaaagaagaaggagaaaccaaggagagtgaagtagtgaagataacagaagaatcaatcctaaaaaggatcaattcaaagaaggaaactaaatctttccaacttgggaaacaactatcatgcaaatggacaacaggtgcaggaccaagaatcggttgtgtaagagattatccatcagagcttcagtttcaagcactggaacaagtgaacttgtctccgagaagtgcttctgtttcaagactctgtttctcttcttcatcgcaaacgcaaacgcctcaaatgtcaccgttatggcgaggaatgtcactacctacagatatcacactcacgaattcataa。
3.如权利要求2所述的拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，其特征在于，所述基因的编码序列的DNA分子来控制拟南芥的开花时间。
4.如权利要求2所述的拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，其特征在于，通过使用所述控制拟南芥的开花时间的基因来控制拟南芥的开花时间。
5.如权利要求2所述的拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，其特征在于，开花时间通过在拟南芥中过度表达拟南芥钙调素结合蛋白基因ATIQM2分子而被加速。
6.如权利要求2所述的拟南芥钙调素结合蛋白基因ATIQM2在拟南芥开花调节中的应用，其特征在于，开花时间通过抑制拟南芥钙调素结合蛋白基因ATIQM2的DNA分子的表达而被延迟。
Priority Applications (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN201410040966.8A CN104098664B (zh) | 2014-01-27 | 2014-01-27 | 拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 |
Applications Claiming Priority (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CN201410040966.8A CN104098664B (zh) | 2014-01-27 | 2014-01-27 | 拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 |
Publications (2)
| Publication Number | Publication Date |
|---|---|
| CN104098664A CN104098664A (zh) | 2014-10-15 |
| CN104098664B true CN104098664B (zh) | 2017-05-10 |
Family
ID=51667240
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| CN201410040966.8A Expired - Fee Related CN104098664B (zh) | 2014-01-27 | 2014-01-27 | 拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 |
Country Status (1)
| Country | Link |
|---|---|
| CN (1) | CN104098664B (zh) |
Families Citing this family (4)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN108004248B (zh) * | 2017-12-07 | 2020-06-12 | 华南农业大学 | 一种黄瓜钙结合蛋白基因CsCaM在提高植物耐热性中的应用 |
| CN110863063B (zh) * | 2019-12-13 | 2022-12-06 | 中国农业科学院深圳农业基因组研究所 | 与黄瓜开花时间性状相关的分子标记 |
| CN116003563A (zh) * | 2021-10-21 | 2023-04-25 | 中国科学院植物研究所 | 钙调素结合蛋白CaMBP13在调控植物耐冷性中的应用 |
| CN114990137B (zh) * | 2022-06-28 | 2023-06-09 | 鲁东大学 | 拟南芥钙结合蛋白基因AtCAREF及应用 |
Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO1997035968A1 (en) * | 1996-03-28 | 1997-10-02 | Washington State University Research Foundation | Compositions and methods for production of male-sterile plants |
-
2014
- 2014-01-27 CN CN201410040966.8A patent/CN104098664B/zh not_active Expired - Fee Related
Patent Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| WO1997035968A1 (en) * | 1996-03-28 | 1997-10-02 | Washington State University Research Foundation | Compositions and methods for production of male-sterile plants |
Non-Patent Citations (2)
| Title |
|---|
| calmodulin-binding family protein [Arabidopsis thaliana];NP_187969.1;《GENBANK》;20140122;1-2 * |
| CML24, Regulated in Expression by Diverse Stimuli, Encodes a Potential Ca2+ Sensor That Functions in Responses to Abscisic Acid, Daylength, and Ion Stress;Nikki A. Delk等;《Plant Physiology》;20050930;第139卷;240-253 * |
Also Published As
| Publication number | Publication date |
|---|---|
| CN104098664A (zh) | 2014-10-15 |
Similar Documents
| Publication | Publication Date | Title |
|---|---|---|
| US20230365987A1 (en) | Transcription factors to improve resistance to environmental stress in plants | |
| Shishkova et al. | Determinate root growth and meristem maintenance in angiosperms | |
| Liu et al. | Phytochrome B control of total leaf area and stomatal density affects drought tolerance in rice | |
| Lellis et al. | Deletion of the eIFiso4G subunit of the Arabidopsis eIFiso4F translation initiation complex impairs health and viability | |
| Li et al. | Developmental, cytological and transcriptional analysis of autotetraploid Arabidopsis | |
| Sun et al. | Chrysanthemum MADS-box transcription factor CmANR1 modulates lateral root development via homo-/heterodimerization to influence auxin accumulation in Arabidopsis | |
| CN111690625B (zh) | 具有除草剂抗性的乙酰乳酸合酶突变蛋白及其应用 | |
| CN104561023B (zh) | 大豆GmCIB1基因和GmCRY2基因及其调控开花及衰老的作用 | |
| Zhuang et al. | Drought inhibition of tillering in Festuca arundinacea associated with axillary bud development and strigolactone signaling | |
| CN101704885A (zh) | 一种控制水稻抽穗期和籽粒大小的蛋白及其编码基因 | |
| CN104098664B (zh) | 拟南芥钙调素结合蛋白基因atiqm2在开花调节中的应用 | |
| CN109576282A (zh) | 月季转录因子RhMYB4及其花器官发育调控中的应用 | |
| Couzigou et al. | The legume NOOT‐BOP‐COCH‐LIKE genes are conserved regulators of abscission, a major agronomical trait in cultivated crops | |
| MULEKE et al. | Genome-wide identification and expression profiling of MYB transcription factor genes in radish (Raphanus sativus L.) | |
| Zhang et al. | Molecular characterization and expression analysis of small heat shock protein 17.3 gene from Sorbus pohuashanensis (Hance) Hedl. in response to abiotic stress | |
| CN111233988B (zh) | 茄子钾离子通道蛋白SmAKT1及其编码基因和应用 | |
| Abdul‐Awal et al. | A sorghum gigantea mutant attenuates florigen gene expression and delays flowering time | |
| Bartlett et al. | Protein change in plant evolution: tracing one thread connecting molecular and phenotypic diversity | |
| Yang et al. | Identification and expression analyzes of CC-type glutaredoxin in cucumber (Cucumis sativus L.) under abiotic stress | |
| CN104278040A (zh) | 一种提高植物抗逆境能力的方法 | |
| CN108997487A (zh) | 抗逆相关蛋白z76在调控植物抗逆性中的应用 | |
| Pi et al. | A GT‐1 and PKc domain‐containing transcription regulator SIMPLE LEAF1 controls compound leaf development in woodland strawberry | |
| Skylar et al. | ELONGATA3 is required for shoot meristem cell cycle progression in Arabidopsis thaliana seedlings | |
| CN114990137B (zh) | 拟南芥钙结合蛋白基因AtCAREF及应用 | |
| Sharma et al. | Osa-miR820 regulatory node primes rice plants to tolerate salt stress in an agronomically advantageous manner |
Legal Events
| Date | Code | Title | Description |
|---|---|---|---|
| C06 | Publication | ||
| PB01 | Publication | ||
| C10 | Entry into substantive examination | ||
| SE01 | Entry into force of request for substantive examination | ||
| GR01 | Patent grant | ||
| GR01 | Patent grant | ||
| CF01 | Termination of patent right due to non-payment of annual fee |
Granted publication date: 20170510 |
|
| CF01 | Termination of patent right due to non-payment of annual fee |