碱基编辑技术及其在作物遗传改良中的应用综述

doi:10.16420/j.issn.0513-353x.2020-0540

摘要/Abstract

摘要：

碱基编辑技术是以CRISPR/Cas系统为基础开发的一种能够对基因组进行定点精准编辑的新技术,包括胞嘧啶碱基编辑系统（cytosine base editor,CBE）,腺嘌呤碱基编辑系统（adenine base editor,ABE）以及引导编辑系统（prime editing,PE）。胞嘧啶碱基编辑系统可以将基因组靶位点处的C/G转换为T/A,腺嘌呤碱基编辑系统可以将靶位点处的A/T转变为G/C,而引导编辑系统则可以实现所有12种类型（C-T、G-A、A-G、T-C、C-A、C-G、G-C、G-T、A-C、A-T、T-A、T-G）碱基的任意替换以及碱基的插入和删除。本文中系统介绍了这3种碱基编辑系统的原理、开发过程、各自的优缺点以及在作物遗传改良中的应用和发展,并展望了碱基编辑技术在农作物育种中的应用前景。

关键词: 碱基编辑技术, 胞嘧啶碱基编辑系统（CBE）, 腺嘌呤剪辑编辑系统（ABE）, 引导编辑系统（PE）, 作物改良

Abstract:

The base editing is a new technology developed on the basis of the CRISPR/Cas system that can precisely edit the genome. It includes cytosine base editor（CBE）,adenine base editor（ABE）and the latest prime editing（PE）. The cytosine base editor can convert C/G to T/A at the target site of the genome,the adenine base editor can convert the A/T to G/C,while the prime editing can realize all 12 types nucleotides replacing（C-T,G-A,A-G,T-C,C-A,C-G,G-C,G-T,A-C,A-T,T-A,T-G）,and the bases insertion and deletion. This article makes a systematic introduction of the principles,development processes,respective advantages and disadvantages of the three base editors,and their applications and developments in crop genetic improvement. Meanwhile,It looks forward to the application prospects of base editing technology in crop breeding.

Key words: base editing, cytosine base editor（CBE）, adenine base editor（ABE）, prime editing（PE）, crop development

中图分类号:

李国斌, 艾国, 韦静, 张俊红. 碱基编辑技术及其在作物遗传改良中的应用综述[J]. 园艺学报, 2021, 48(4): 719-732.

LI Guobin, AI Guo, WEI Jing, ZHANG Junhong. Review of Base Editing Technology and its Application in Crop Genetic Improvement[J]. Acta Horticulturae Sinica, 2021, 48(4): 719-732.

图/表 3

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物种 Crop name	编辑系统 Base editor	编辑形式 Editor type	PAM	靶基因 Targeted gene	应用 Function	参考文献 Reference
水稻Rice	BE3, Anc689BE4max	C-T	NGG,NG	NRT1.1b,SLR1	氮利用率、株形 Nitrogen use efficiency,Plant architecture	Lu & Zhu, 2017;Wang et al.2019
	CBE-P1-P5, ABE-P1-P5	C-T,A-G	NGG	OsSPL13, OsSPL14, OsSPL16, OsSPL17, OsSPL18, GRF4, OsTOE1, OslDS1, OMTN1, SNB, PMS1, PMS3	株形,产量,抗性 Plant architecture,Yield increase,Improve resistance	Hua et al.2019
	ABE-P1S, ABEmax	A-G	NG,NGG	OsSPL14, OsSPL16, OsSERK2, SLR1, Tms9-1, OsNRT1.1B, OsACC1, OsDEP1, OsSPL17, OsSPL18, SLR1, ALS, EPSPS,	株形,抗除草剂 Plant architecture,Herbicide resistance	Hua et al.2018,2020b;Wang et al.2019
	rBE3,rBE4, rBE5,rBE9	C-T	NGG, NAG, NGA, AGTG, AGCG AGTG, AGCG	OsSERK1, OsSERK2, OsAOS1, OsJAR1, OsJAR2, OsCOL2		Ren et al.2017,2018
	rBE14	A-G	NGG	OsMPK6, OsMPK13, OsSERK2, OsWRKY45, Tms9-1	抗病Pathogen resistance	Yan et al.2018
	rBE22, rBE25	C-T	NG, NAC, NTG, NTT, NCG	OsBZR1		Ren et al.2019;Zhong et al.2019;Wang et al.2020
	rBE23, rBE26	A-G	NG, NAC, NTG, NTT, NCG	OsCERK2, OsGS1	抗病 Pathogen resistance	Ren et al.2019;Wang et al.2020
	pCXUN-BE3	C-T	NGG	OsPDS, OsSBEIIb	品质 Nutritional improvement	Li et al.2017
	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	OsAAT-T1, OsCDC48, OsDEP1, sNRT1.1B-T1, OsOD, OsEV		Zong et al.2018
	xCas9-epBE	C-T	GA, NG	OsMPK2, OsMPK5, OsALS, OsNRT1.1B, OsWaxy	抗病, 抗除草剂 Pathogen resistance, Herbicide resistance	Zhong et al.2019;Zhang et al.2020
小麦Wheat	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	TaALS, TaMTL, TaLOX2-T1, TaLOX2-T2	抗除草剂 Herbicide resistance	Zong et al.2018
玉米Maize	PBE（BE3）	C-T	NGG	ZmCENH3		Zong et al.2017
马铃薯 Potato	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	StGBSS-T6		Zong et al.2018
	CBE	C-T	NGG	StGBSS1	淀粉含量 Starch synthesis	Veillet et al.2019a
拟南芥 Arabidopsis	BE3	C-T	NGG	AtALS	抗除草剂 Herbicide resistance	Chen et al.2017
番茄Tomato	CBE	C-T	NGG	ALS	抗除草剂 Herbicide resistance	Veillet et al.2019b
	Target-AID	C-T	NGG	DELLA, ETR1	株型 Plant architecture	Shimatani et al.2017
油菜Rape	pcABE7.10	A-G	NGG	BnALS, BnPDS	抗除草剂 Herbicide resistance	Kang et al.2018
	CBE	C-T	NGG	BnALS1, BnALS3	抗除草剂 Herbicide resistance	Wu et al.2020
棉花Cotton	GhBE3	C-T	NGG	GhCAL, GhPEPB	株型 Plant architecture	Qin et al.2020
西瓜Watermelon	BE3	C-T	NGG	ClALS	抗除草剂 Herbicide resistance	Tian et al.2018

物种 Crop name	编辑系统 Base editor	编辑形式 Editor type	PAM	靶基因 Targeted gene	应用 Function	参考文献 Reference
水稻Rice	BE3, Anc689BE4max	C-T	NGG,NG	NRT1.1b,SLR1	氮利用率、株形 Nitrogen use efficiency,Plant architecture	Lu & Zhu, 2017;Wang et al.2019
	CBE-P1-P5, ABE-P1-P5	C-T,A-G	NGG	OsSPL13, OsSPL14, OsSPL16, OsSPL17, OsSPL18, GRF4, OsTOE1, OslDS1, OMTN1, SNB, PMS1, PMS3	株形,产量,抗性 Plant architecture,Yield increase,Improve resistance	Hua et al.2019
	ABE-P1S, ABEmax	A-G	NG,NGG	OsSPL14, OsSPL16, OsSERK2, SLR1, Tms9-1, OsNRT1.1B, OsACC1, OsDEP1, OsSPL17, OsSPL18, SLR1, ALS, EPSPS,	株形,抗除草剂 Plant architecture,Herbicide resistance	Hua et al.2018,2020b;Wang et al.2019
	rBE3,rBE4, rBE5,rBE9	C-T	NGG, NAG, NGA, AGTG, AGCG AGTG, AGCG	OsSERK1, OsSERK2, OsAOS1, OsJAR1, OsJAR2, OsCOL2		Ren et al.2017,2018
	rBE14	A-G	NGG	OsMPK6, OsMPK13, OsSERK2, OsWRKY45, Tms9-1	抗病Pathogen resistance	Yan et al.2018
	rBE22, rBE25	C-T	NG, NAC, NTG, NTT, NCG	OsBZR1		Ren et al.2019;Zhong et al.2019;Wang et al.2020
	rBE23, rBE26	A-G	NG, NAC, NTG, NTT, NCG	OsCERK2, OsGS1	抗病 Pathogen resistance	Ren et al.2019;Wang et al.2020
	pCXUN-BE3	C-T	NGG	OsPDS, OsSBEIIb	品质 Nutritional improvement	Li et al.2017
	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	OsAAT-T1, OsCDC48, OsDEP1, sNRT1.1B-T1, OsOD, OsEV		Zong et al.2018
	xCas9-epBE	C-T	GA, NG	OsMPK2, OsMPK5, OsALS, OsNRT1.1B, OsWaxy	抗病, 抗除草剂 Pathogen resistance, Herbicide resistance	Zhong et al.2019;Zhang et al.2020
小麦Wheat	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	TaALS, TaMTL, TaLOX2-T1, TaLOX2-T2	抗除草剂 Herbicide resistance	Zong et al.2018
玉米Maize	PBE（BE3）	C-T	NGG	ZmCENH3		Zong et al.2017
马铃薯 Potato	A3A-PBE	C-T	NGG, NGA, NGCG, NNGRRT, NNNRRT	StGBSS-T6		Zong et al.2018
	CBE	C-T	NGG	StGBSS1	淀粉含量 Starch synthesis	Veillet et al.2019a
拟南芥 Arabidopsis	BE3	C-T	NGG	AtALS	抗除草剂 Herbicide resistance	Chen et al.2017
番茄Tomato	CBE	C-T	NGG	ALS	抗除草剂 Herbicide resistance	Veillet et al.2019b
	Target-AID	C-T	NGG	DELLA, ETR1	株型 Plant architecture	Shimatani et al.2017
油菜Rape	pcABE7.10	A-G	NGG	BnALS, BnPDS	抗除草剂 Herbicide resistance	Kang et al.2018
	CBE	C-T	NGG	BnALS1, BnALS3	抗除草剂 Herbicide resistance	Wu et al.2020
棉花Cotton	GhBE3	C-T	NGG	GhCAL, GhPEPB	株型 Plant architecture	Qin et al.2020
西瓜Watermelon	BE3	C-T	NGG	ClALS	抗除草剂 Herbicide resistance	Tian et al.2018