Article(id=1220655368200437804, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1220655362521351042, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2020-0942, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1591632000000, receivedDateStr=2020-06-09, revisedDate=1593014400000, revisedDateStr=2020-06-25, acceptedDate=null, acceptedDateStr=null, onlineDate=1768956518459, onlineDateStr=2026-01-21, pubDate=1594483200000, pubDateStr=2020-07-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1768956518459, onlineIssueDateStr=2026-01-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1768956518459, creator=13701087609, updateTime=1768956518459, updator=13701087609, issue=Issue{id=1220655362521351042, tenantId=1146029695717560320, journalId=1189982191388893191, year='2020', volume='55', issue='7', pageStart='1357', pageEnd='1706', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1768956517105, creator=13701087609, updateTime=1768957228011, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1220658344335950505, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1220655362521351042, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1220658344335950506, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1220655362521351042, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1562, endPage=1572, ext={EN=ArticleExt(id=1220655368821194821, articleId=1220655368200437804, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Application of base editors in the treatment of genetic disorders, columnId=1190335348648547107, journalTitle=Acta Pharmaceutica Sinica, columnName=Reviews, runingTitle=null, highlight=null, articleAbstract=
To date, CRISPR/Cas systems represent the most widely used tool for genome editing; however, its application scope for gene therapy has been largely limited due to its limited efficiency in activating homologydirected repair for DNA and off-target effect. Base editing is a new CRISPR/Cas-based genome-editing strategy, which allows single nucleotide to be precisely corrected in a narrow window scope on the target DNA or RNA by taking advantage of different nucleobase deaminases. Base editors include cytosine base editors (CBEs) and adenine base editors (ABEs), which can induce the conversions from C·G to T·A and A·T to G·C, respectively. Base editors work independently of double-strand DNA breaks (DSBs) and DNA donor templates, and thus they are extensively adopted for a wide range of therapeutic applications for genetic diseases, largely owing to their high efficiency and great specificity. In this review, we summarize the development of base editors and their potentials as therapeutic drugs for treating genetic diseases, and future outlooks are also discussed.
, correspAuthors=Yuan PING, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2020 Acta Pharmaceutica Sinica. All rights reserved., copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Yuan CHEN, Yu-xuan CHEN, Yuan PING), CN=ArticleExt(id=1220655370696048809, articleId=1220655368200437804, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=碱基编辑系统在遗传性疾病治疗中的应用, columnId=1190335349655180086, journalTitle=药学学报, columnName=综述, runingTitle=null, highlight=null, articleAbstract=
CRISPR/Cas系统是现阶段应用最广泛的基因组编辑工具，但由于其激活同源重组进行DNA修复的效率低且存在脱靶效应，严重限制了该工具在基因治疗方面的应用。碱基编辑是一种基于CRISPR/Cas系统的基因编辑新策略，其工作原理是利用其结合的碱基脱氨酶精确编辑目标DNA或RNA上一小段窗口内的核苷酸。碱基编辑系统包括胞嘧啶碱基编辑系统（CBEs）和腺嘌呤碱基编辑系统（ABEs），可以分别实现碱基对C·G到T·A和A·T到G·C的转换。碱基编辑系统作用过程中不依赖于DNA双链断裂的发生，无需引入供体DNA，并因其高效性及特异性已被广泛应用于遗传性疾病治疗的研究中。本文综述了近年来碱基编辑系统的发展历程和在遗传性疾病治疗中的潜力，并对其作为基因治疗药物的发展前景进行了展望。
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10.1016/j.cell.2020.05.037., articleTitle=Determinants of base editing outcomes from target library analysis and machine learning, refAbstract=null)], funds=[Fund(id=1220655374974239289, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, awardId=81872807, language=CN, fundingSource=国家自然科学基金面上项目(81872807), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1220655370968678587, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, xref=null, ext=[AuthorCompanyExt(id=1220655370972872892, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, companyId=1220655370968678587, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=College of Pharmaceutical Sciences, Zhejiang University, Hangzhou 310058, China), AuthorCompanyExt(id=1220655370981261501, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, companyId=1220655370968678587, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=浙江大学药学院, 浙江 杭州 310058)])], figs=[ArticleFig(id=1220655373124551067, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=TWhEkt8OK8FRQgStCSKMHQ==, figureFileBig=dQfMv+bUVNFLAfTiJLvjiQ==, tableContent=null), ArticleFig(id=1220655373233602980, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 1, caption=
Working mechanisms of cytosine base editors (BE1, BE2, BE3 or BE4). a: Chemical reaction of cytosine (C) to thymidine (T) conversion. C deamination generates uridine (U), which base pairs as T. R represents 2'-deoxyribose in DNA; b: Schematic illustration of working mechanisms of cytosine base editors. Adapted from Ref. 4 with permission. Copyright © 2018 Springer Nature , figureFileSmall=TWhEkt8OK8FRQgStCSKMHQ==, figureFileBig=dQfMv+bUVNFLAfTiJLvjiQ==, tableContent=null), ArticleFig(id=1220655373464289716, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=MUP8cQII1Hv6dRdRx8GRbQ==, figureFileBig=9nma4Y6LaOxr2qIUKyq5MQ==, tableContent=null), ArticleFig(id=1220655373577535934, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 2, caption=
Working mechanisms of adenine base editors. a: Chemical reaction of adenine (A) to guanine (G) conversion. R stands for 2'-deoxyribose in DNA or ribose in RNA; b: Schematic illustration of working mechanisms of adenine base editors. Adapted from Ref. 4 with permission. Copyright © 2018 Springer Nature , figureFileSmall=MUP8cQII1Hv6dRdRx8GRbQ==, figureFileBig=9nma4Y6LaOxr2qIUKyq5MQ==, tableContent=null), ArticleFig(id=1220655373665616324, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=OKo/OaNiZDa3ZGqkqoLhWg==, figureFileBig=l1YesoH93+as3nU8wc7/IQ==, tableContent=null), ArticleFig(id=1220655373766279631, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 3, caption=
Schematic illustration of the correction of the mutated <i>Pah<sup>enu2</sup></i> locus in cell lines, liver organoids and <i>Pah<sup>enu2</sup></i> mice. Adapted from Ref. 16 with permission. Copyright © 2018 Springer Nature , figureFileSmall=OKo/OaNiZDa3ZGqkqoLhWg==, figureFileBig=l1YesoH93+as3nU8wc7/IQ==, tableContent=null), ArticleFig(id=1220655373862748632, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=te8TuNVYeRhjeHZNifxfmw==, figureFileBig=qZUBXIcIZ9vnnymT1kVxcQ==, tableContent=null), ArticleFig(id=1220655373963411939, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 4, caption=
Application of adenine base editing for the treatment of mouse model of tyrosinaemia. a: Exon skipping caused by G > A mutation (red) at the last nucleotide of exon 8 in the <i>Fah</i> gene of <i>Fah<sup>mut/mut</sup></i> mice. Exon sequences are shown in upper case. The G > A mutation is at position 9 of the sgRNA target. WT: Wild-type allele; b: Schematic illustration of hydrodynamic injection of adenine base editors (ABEs) plasmids for the treatment of <i>Fah<sup>mut/mut</sup></i> mice. Adapted from Ref. 19 with permission. Copyright © 2020 Springer Nature , figureFileSmall=te8TuNVYeRhjeHZNifxfmw==, figureFileBig=qZUBXIcIZ9vnnymT1kVxcQ==, tableContent=null), ArticleFig(id=1220655374055686634, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=N/HOeHEqijR70tyj8ADDbw==, figureFileBig=Wg9HpYna+snMYHhRWVcm3w==, tableContent=null), ArticleFig(id=1220655374131184114, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 5, caption=
Schematic illustration of <i>HBB-28</i> (A > G) mutation and potential therapeutic base editing by targeting Cs at -28 and -25 in the editing window with an <i>HBB</i>-targeting gRNA. Mutations to the bystander cytidine at the -25 position are deleterious and cause <i>β</i>-thalassemia phenotypes independent of the identity of the -28 nucleotide. Adapted from Ref. 24 with permission. Copyright © 2018 Springer Nature , figureFileSmall=N/HOeHEqijR70tyj8ADDbw==, figureFileBig=Wg9HpYna+snMYHhRWVcm3w==, tableContent=null), ArticleFig(id=1220655374290567674, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=Ae09w+Id9JL0uABHHAcPoA==, figureFileBig=JLXg/ynS3C7j+FWF+O6s1w==, tableContent=null), ArticleFig(id=1220655374399619587, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Figure 6, caption=
Application of ABEs for the treatment of a mouse model of Duchenne muscular dystrophy. a: The ABEs target sequence in exon 20 of the <i>Dmd</i> gene contains a nonsense mutation. The PAM sequence is shown in blue, and the protospacer sequence is underlined. The mutated nucleotide in the <i>Dmd</i> knockout mouse and the guanine nucleotide corrected by ABEs are shown in red and green, respectively; b: Schematic diagram of <i>trans</i>-splicing AAV vectors encoding engineered <i>E. coli</i> TadA (ecTadA), which are conjugated to the first half of nickase Cas9 (nCas9-NT) and the second half of nickase Cas9 (nCas9-CT). SD: Splicing donor; SA: Splicing acceptor. Adapted from Ref. 29 with permission. Copyright © 2018 Springer Nature , figureFileSmall=Ae09w+Id9JL0uABHHAcPoA==, figureFileBig=JLXg/ynS3C7j+FWF+O6s1w==, tableContent=null), ArticleFig(id=1220655374512865805, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
| Item | Base editing system | Homology directed repair |
| DSB | Unnecessary | Necessary |
| Donor DNA | Unnecessary | Necessary |
| Cell cycle | Unrestricted | Restricted to G2 and S phases of the cell cycle |
| Indels | Lower | Higher |
| Editing efficiency in vitro and in vivo | Higher | Lower |
), ArticleFig(id=1220655374638694934, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Table 1, caption=
Comparison of base editing system and homology directed repair. DSB: Double-strand DNA break; Indels: Insertion and deletion
, figureFileSmall=null, figureFileBig=null, tableContent=
| Item | Base editing system | Homology directed repair |
| DSB | Unnecessary | Necessary |
| Donor DNA | Unnecessary | Necessary |
| Cell cycle | Unrestricted | Restricted to G2 and S phases of the cell cycle |
| Indels | Lower | Higher |
| Editing efficiency in vitro and in vivo | Higher | Lower |
), ArticleFig(id=1220655374764524066, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
| Cytosine base editor | Base editor architecture | Editing window | PAM | Reference |
| BE1 | rAPOBEC1-Sp dCas9 | 4 - 8 | NGG | [6] |
| BE2 | rAPOBEC1-Sp dCas9-UGI | 4 - 8 | NGG | [6] |
| BE3 | rAPOBEC1-Sp nCas9 (D10A)-UGI | 4 - 8 | NGG | [6] |
| Target-AID | Sp nCas9 (D10A)-PmCDA1-UGI | 2 - 8 | NGG | [8] |
| CRISPR-X | Sp dCas9; MS2-hAIDΔ | -50 - 50 | NGG | [9] |
| TAM | Sp dCas9-hAID*-P182X | 4 - 10 | NGG | [10] |
| BE4 | rAPOBEC1-Sp nCas9 (D10A)-UGI-UGI | 4 - 8 | NGG | [11] |
| BE4-Gam | Gam-rAPOBEC1-Sp nCas9 (D10A)-UGI-UGI | 4 - 8 | NGG | [11] |
), ArticleFig(id=1220655374860993071, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1220655368200437804, language=CN, label=Table 2, caption=
Cytosine base editors with various characteristics. PAM: Protospacer adjacent motif; BE: Base editor; Target-AID: Target activation-induced cytidine deaminase; TAM: Targeted AID-mediated mutagenesis
, figureFileSmall=null, figureFileBig=null, tableContent=
| Cytosine base editor | Base editor architecture | Editing window | PAM | Reference |
| BE1 | rAPOBEC1-Sp dCas9 | 4 - 8 | NGG | [6] |
| BE2 | rAPOBEC1-Sp dCas9-UGI | 4 - 8 | NGG | [6] |
| BE3 | rAPOBEC1-Sp nCas9 (D10A)-UGI | 4 - 8 | NGG | [6] |
| Target-AID | Sp nCas9 (D10A)-PmCDA1-UGI | 2 - 8 | NGG | [8] |
| CRISPR-X | Sp dCas9; MS2-hAIDΔ | -50 - 50 | NGG | [9] |
| TAM | Sp dCas9-hAID*-P182X | 4 - 10 | NGG | [10] |
| BE4 | rAPOBEC1-Sp nCas9 (D10A)-UGI-UGI | 4 - 8 | NGG | [11] |
| BE4-Gam | Gam-rAPOBEC1-Sp nCas9 (D10A)-UGI-UGI | 4 - 8 | NGG | [11] |
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