碱基编辑系统研究进展

宗媛,高彩霞

Progress on base editing systems

Zong Yuan,Gao Caixia

表2 各种碱基编辑器(BE)的特点

Table 2 Base editors with various characteristics

BE名称 脱氨酶 Cas蛋白 PAM 类型 UGI 个数 NLS 个数 密码子 优化 编辑 活性窗口 参考 文献 BE1 rAPOBEC1 dCas9 NGG - 1× - 4~8 [18] BE2 rAPOBEC1 dCas9 NGG 1× 1× - 4~8 [18] BE3 rAPOBEC1 nCas9 NGG 1× 1× - 4~8 [18] PBE rAPOBEC1 nCas9 NGG 1× 3× + 3~9 [82] HF-BE3 rAPOBEC1 HF1-nCas9 NGG 1× 1× - 4~8 [123] HF2-BE2 rAPOBEC1 HF2-dCas9 NGG 1× 1× - 4~8 [58] BE4 rAPOBEC1 nCas9 NGG 2× 1× - 4~8 [23] BE4-Gam rAPOBEC1 nCas9 NGG 2× 1× - 4~8 [23] BE4max rAPOBEC1 nCas9 NGG 2× 2× - 4~8 [26] AncBE4max rAPOBEC1 nCas9 NGG 2× 2× + 4~8 [26] eBE-S3 rAPOBEC1 nCas9 NGG 4× 1× - 4~8 [24] FNLS-BE3 rAPOBEC1 nCas9 NGG 1× 2× + 3~8 [27] VQR-BE3 rAPOBEC1 nVQRCas9 NGA 1× 1× - 4~11 [29] EQR-BE3 rAPOBEC1 nEQRCas9 NGAG 1× 1× - 4~11 [29] VRER-BE3 rAPOBEC1 nVRERCas9 NGCG 1× 1× - 3~10 [29] SaBE3 rAPOBEC1 nSaCas9 NNGRRT 1× 1× - 3~12 [29] SaBE4 rAPOBEC1 nSaCas9 NNGRRT 2× 1× - 3~12 [23] SaBE4-Gam rAPOBEC1 nSaCas9 NNGRRT 2× 1× - 3~12 [23] SaKKH-BE3 rAPOBEC1 nSaKKHCas9 NNNRRT 1× 1× - 3~12 [29] Spy-macnCas9-BE3 rAPOBEC1 Spy-nmacCas9 NAA 1× 1× - 4~8 [31] dCpf1-eBE rAPOBEC1 dCpf1 TTTV 4× 3× - 8~13 [32] enAsBE1.1-1.4 rAPOBEC1 denAsCas12a TTTV 1× 1×-4× - -5~25 [33] xBE3 rAPOBEC1 nxCas9 NG 1× 1× - 4~8 [34] BE3-NG rAPOBEC1 nSpCas9-NG NG 1× 1× - 4~8 [35] YE1-BE3 rAPOBEC1-YE1 nCas9 NGG 1× 1× - 4~7 [29] YE2-BE3 rAPOBEC1-YE2 nCas9 NGG 1× 1× - 5~6 [29] EE-BE3 rAPOBEC1-EE nCas9 NGG 1× 1× - 5~6 [29] YEE-BE3 rAPOBEC1-YEE nCas9 NGG 1× 1× - 5~6 [29] BE3-PAPAPAP rAPOBEC1 nCas9 NGG 1× 1× - 5~6 [36] BE3-R33A rAPOBEC1-R33A nCas9 NGG 1× 1× - 5~7 [131] BE3-R33A/K34A rAPOBEC1-R33A/K34A nCas9 NGG 1× 1× - 5~6 [131] BE-PLUS rAPOBEC1-scFV nCas9 NGG 1× 1× - 4~16 [37] CP-CBEmax rAPOBEC1 CP-nCas9 NGG 2× 1× - 4~11 [25] eA3A-BE3 hAPOBEC3A-N57G nCas9 NGG 1× 1× - 4~8 [41] eA3A-HF1-BE3-2xUGI hAPOBEC3A-N57G HF1-nCas9 NGG 2× 1× - 4~8 [41] eA3A-Hypa-BE3-2xUGI hAPOBEC3A-N57G nHypaCas9 NGG 2× 1× - 4~8 [41] hA3A-BE3 hAPOBEC3A nCas9 NGG 1× 1× - 2~13 [38] hA3A-BE3-Y130F hAPOBEC3A-Y130F nCas9 NGG 1× 1× - 3~8 [38] A3A-PBE hAPOBEC3A nCas9 NGG 1× 2× + 1~17 [28] DBE-A3A hAPOBEC3A-MS2 nCas9 NGG 1× 1× - 2~17 [39] hA3AR128A hAPOBEC3A-R128A nCas9 NGG 1× 1× - 3~9 [133] A3G-BE4max hAPOBEC3G-CTD nCas9 NGG 2× 1× - 4~10 [42] Target-AID PmCDA1 nCas9 NGG 1× 1× - 2~8 [19] Target-AID-NG PmCDA1 nCas9 NG 1× 1× - 2~8 [35] nCDA1-BE3 PmCDA1Δ nCas9 NGG 1× 1× - 3~4 [36] TAM hAIDx dCas9 NGG 1× 1× - 4~10 [20] CRISPR-X hAIDΔ-MS2 dCas9 NGG - 1× - -50~50 [21] eAID-BE4max hAID nCas9 NGG 2× 2× + 4~8 [40] DBE-AIDmono hAID-mono-MS2 nCas9 NGG 1× 1× - 2~17 [39] ZF-AID hAID ZF - 1× 1× - - [22] TALE-AID hAID TALE - 1× 1× - - [22] ABE7.9 TadA-TadA* nCas9 NGG 1× 1× - 4~9 [107] ABE7.10 TadA-TadA* nCas9 NGG - 1× - 4~7 [107] PABE-7 TadA-TadA* nCas9 NGG - 1× + 4~7 [108] ABEmax TadA-TadA* nCas9 NGG - 2× + 4~7 [26] VQR-ABE TadA-TadA* nVQRCas9 NGA - 1× +/- 4~8 [30,114] VQR-ABEmax TadA-TadA* nVQRCas9 NGA - 2× + 4~8 [25] VRER-ABE TadA-TadA* nVRERCas9 NGCG - 1× + 4~8 [30] VRER-ABEmax TadA-TadA* nVRERCas9 NGCG - 2× + 4~8 [25] VRQR-ABEmax TadA-TadA* nVRQRCas9 NGA - 2× + 4~8 [25] ABEsa TadA-TadA* nSaCas9 NNGRRT - 1× + 6~12 [111] SaABEmax TadA-TadA* nSaCas9 NNGRRT - 2× + 4~14 [25] SaKKH-ABE TadA-TadA* nSaKKHCas9 NNNRRT - 1× + 8~13 [30,117] SaKKH-ABEmax TadA-TadA* nSaKKHCas9 NNNRRT - 2× + 4~14 [25] xABE TadA-TadA* nxCas9 NG - 1× - 4~7 [34] xABEmax TadA-TadA* nxCas9 NG - 2× + 4~8 [25] ABE-NG TadA-TadA* nSpCas9-NG NG - 1× + 4~7 [74] ABE-NG-S TadA* nSpCas9-NG NG - 1× + 4~7 [74] NG-ABEmax TadA-TadA* nSpCas9-NG NG - 2× - 4~8 [35] CP-ABEmax TadA-TadA* CP-nCas9 NGG - 2× + 4~12 [25] ScCas9-ABE(7.10) TadA-TadA* nScCas9 NNGN - 1× - 4~7 [109] ABEmaxAW TadA-E59A-TadA*-V106W nCas9 NGG - 2× + 4~8 [132] ABEmaxQW TadA-E59Q-TadA*-V106W nCas9 NGG - 2× + 4~8 [132] ABE7.10F148A TadA-F148A-TadA*-F148A nCas9 NGG - 1× - 5 [133]