Advance in Gene-editing Technology Based on CRISPR/Cas9 and Its Application in Plants

doi:10.11983/CBB22020

Abstract

Abstract: CRISPR/Cas9 technology is a gene editing strategy using Cas9 nuclease guided by RNA to target an interest gene in genome. Recently, a large number of new-type gene editors based on the CRISPR/Cas9 have been updated rapidly and become more precisely and efficiently in gene editing, which has a great application prospect in crop molecular-designing breeding. This paper summarized the technical principles, editing effects and applications of the CRISPR/cas9 and its related gene editors, and also discussed the aspects of dilemmas, countermeasures and prospects, intending to provide reference for scientific researchers in the related fields.

Key words: CRISPR/Cas9, gene editing, technical progress, application in plants

He Xiaoling, Liu Pengcheng, Ma Bojun, Chen Xifeng. Advance in Gene-editing Technology Based on CRISPR/Cas9 and Its Application in Plants[J]. Chinese Bulletin of Botany, 2022, 57(4): 508-531.

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URL: https://www.chinbullbotany.com/EN/10.11983/CBB22020

https://www.chinbullbotany.com/EN/Y2022/V57/I4/508

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References 110

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编辑技术	编辑器	脱氨酶/逆转录酶		Cas蛋白	PAM (5′-3′)		活性窗口	编辑类型		特点	参考文献
胞嘧啶碱基编辑(CBEs)	BE3	rAPOBEC1		nCas9^D10A	NGG		C₄-C₈	C:G>T:A		实现C>T替换, 无需DSB或提供模板	Komor et al., 2016
	nCas9-PBE	rAPOBEC1		nCas9^D10A	NGG		C₃-C₉			在植物中实现精准高效的C>T替换	Zong et al., 2017
	Target-AID	PmCDA1		nCas9^D10A	NGG		C₁-C₅			具有较好的GC碱基编辑能力	Nishida et al., 2016
	dCas9-AIDx	hAID		dCas9	NCN		C₄-C₉			具有较好的GC碱基编辑能力	Ma et al., 2016
	A3A^Y130F- CBE_V01 A3A^Y130F- CBE_V04	hAPOBEC3A^Y130F		nCas9^D10A	NGG		C₄-C₁₅			活性窗口拓宽至12 nt	Ren et al., 2021a
	eCDAL	LjCDA1L-4		nCas9^D10A	NGG		C₁-C₁₂			活性窗口拓宽至12 nt	Xu et al., 2021
	BE3-PAPAPAP	rAPOBEC1		nCas9^D10A	NGG		C₅-C₆			活性窗口缩小至1-2 nt	Tan et al., 2019
	PhieCBEs	evorAPOCBEC1 evoFERNY evoCDA1 hA3A		nCas9-NG eCas9n- NG	NG NGG		C_-1-C₁₅			编辑窗口广	Zeng et al., 2020
	BE3^R126E BE3^R132E YE1-BE3	rAPOBEC1^R126E rAPOBEC1^R132E rAPOBEC1^W90Y+R126E		nCas9^D10A	NGG		C₅-C₇			脱靶率显著降低	Zuo et al., 2020
腺嘌呤碱基编辑(ABEs)	ABE7.10	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₄-A₇	A:T>G:C		实现A>G替换, 无需DSB或提供模板	Gaudelli et al., 2017
	PABE	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₄-A₈			在植物中实现精准高效的A>G替换	Li et al., 2018b
	ABE-Ps	ecTadA:ecTadA^*		nCas9^D10A SaCas9	NGG NNNRRT		A₁-A₁₅			在植物中实现精准高效的A>G替换	Hua et al., 2018
	rBE14	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₅-A₇			荧光检测编辑植株	Yan et al., 2018
	rABE8e	TadA8e^V106W		nCas9^D10A nCas9-NG	NG NGG		A₅-A₆			极高的靶点编辑效率和碱基纯合替换效率	Wei et al., 2021
	PhieABEs	TadA8e		nCas9-NG SpGn SpRYn	NG NGN NNN		A₁-A₁₄			编辑效率高, 几近无PAM靶向, 窗口广	Tan et al., 2022
	SpRY-ABE8e	ecTadA:ecTadA^*		nCas9^D10A	NNN		A₃-A₁₀			几近无PAM靶向	Ren et al., 2021b
	ABE7.10^F148A	ecTadA^F148A: ecTadA^*F148A		nCas9^D10A	NGG		A₅-A₆			活性窗口缩小至1-2 nt	Zhou et al., 2019
CG碱基编辑(GBEs)	GBE	AID rAPOBEC1		nCas9^D10A	NGG		C₃-C₇	C:G>A:T/G:C		实现嘧啶与嘌呤间的颠换	Zhao et al., 2021
CG碱基编辑(GBEs)	CGBE	rAPOBEC1		nCas9^D10A	NGG		C₅-C₆	C:G>A:T/G:C		实现嘧啶与嘌呤间的颠换	Chen et al., 2021
双碱基编辑(A&CBE)	A&C-BEmax	hAID ecTadA:ecTadA^*		nCas9^D10A	NGG		C₂-C₁₇ A₄-A₇	C:G>T:A A:T>G:C		实现A、C共编辑	Zhang et al., 2020
	Target-ACE	PmCDA1 ecTadA:ecTadA^*			NGG		C₁-C₁₀ A₄-A₈				Sakata et al., 2019
	SPACE	PmCDA1 ecTadA^*			NGG		C₂-C₇ A₄-A₇				Grünewald et al., 2020
	ACBE	PmCDA1 ecTadA:ecTadA^*			NGG		C₁-C₇ A₄-A₆				Xie et al., 2020
	STEME	hAPOBEC3A ecTadA:ecTadA^*			NG NGD		C₁-C₁₇ A₄-A₈			在植物中实现A、 C共编辑	Li et al., 2020a
	pDuBE1	TadA8e			NGG		C₁-C₁₀ A₂-A₉			在植物中实现A、 C共编辑	Xu et al., 2021
编辑技术	编辑器		脱氨酶/逆转录酶	Cas蛋白	PAM (5′-3′)	活性窗口			编辑类型	特点	参考文献
先导编辑(PEs)	PE		M-MLV RT	nCas9^H840A	NGG	1-50			12种碱基替换插入(<15 bp) 缺失(<40 bp)	不受PAM的距离限制	Anzalone et al., 2019
	PPE		CaMV RT Retron RT							在植物中实现PE 应用	Lin et al., 2020
	pPE2		M-MLV RT							在植物中实现PE 应用	Xu et al., 2020a
	ePPE		M-MLV RT^{∆RNase H} M-MLV RT:NC			1-92			12种碱基替换插入(<40 bp) 缺失(<100 bp)	提高了在植物中的编辑效率	Zong et al., 2022
多重编辑	SWISSs		rAPOBEC1 ecTadA:ecTadA^*	nCas9^D10A	NG	C₃-C₁₆ A₄-A₇			C:G>T:A A:T>G:C Indels	具有A>G、C>T和Indels三重编辑功能	Li et al., 2020b
片段删除编辑	AFIDs		hAPOBEC3A hAPOBEC3Bctd	Cas9	NGG	C₁-C₁₄			多核苷酸删除	精准高效、可预测的多核苷酸缺失	Wang et al., 2020

编辑技术	编辑器	脱氨酶/逆转录酶		Cas蛋白	PAM (5′-3′)		活性窗口	编辑类型		特点	参考文献
胞嘧啶碱基编辑(CBEs)	BE3	rAPOBEC1		nCas9^D10A	NGG		C₄-C₈	C:G>T:A		实现C>T替换, 无需DSB或提供模板	Komor et al., 2016
	nCas9-PBE	rAPOBEC1		nCas9^D10A	NGG		C₃-C₉			在植物中实现精准高效的C>T替换	Zong et al., 2017
	Target-AID	PmCDA1		nCas9^D10A	NGG		C₁-C₅			具有较好的GC碱基编辑能力	Nishida et al., 2016
	dCas9-AIDx	hAID		dCas9	NCN		C₄-C₉			具有较好的GC碱基编辑能力	Ma et al., 2016
	A3A^Y130F- CBE_V01 A3A^Y130F- CBE_V04	hAPOBEC3A^Y130F		nCas9^D10A	NGG		C₄-C₁₅			活性窗口拓宽至12 nt	Ren et al., 2021a
	eCDAL	LjCDA1L-4		nCas9^D10A	NGG		C₁-C₁₂			活性窗口拓宽至12 nt	Xu et al., 2021
	BE3-PAPAPAP	rAPOBEC1		nCas9^D10A	NGG		C₅-C₆			活性窗口缩小至1-2 nt	Tan et al., 2019
	PhieCBEs	evorAPOCBEC1 evoFERNY evoCDA1 hA3A		nCas9-NG eCas9n- NG	NG NGG		C_-1-C₁₅			编辑窗口广	Zeng et al., 2020
	BE3^R126E BE3^R132E YE1-BE3	rAPOBEC1^R126E rAPOBEC1^R132E rAPOBEC1^W90Y+R126E		nCas9^D10A	NGG		C₅-C₇			脱靶率显著降低	Zuo et al., 2020
腺嘌呤碱基编辑(ABEs)	ABE7.10	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₄-A₇	A:T>G:C		实现A>G替换, 无需DSB或提供模板	Gaudelli et al., 2017
	PABE	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₄-A₈			在植物中实现精准高效的A>G替换	Li et al., 2018b
	ABE-Ps	ecTadA:ecTadA^*		nCas9^D10A SaCas9	NGG NNNRRT		A₁-A₁₅			在植物中实现精准高效的A>G替换	Hua et al., 2018
	rBE14	ecTadA:ecTadA^*		nCas9^D10A	NGG		A₅-A₇			荧光检测编辑植株	Yan et al., 2018
	rABE8e	TadA8e^V106W		nCas9^D10A nCas9-NG	NG NGG		A₅-A₆			极高的靶点编辑效率和碱基纯合替换效率	Wei et al., 2021
	PhieABEs	TadA8e		nCas9-NG SpGn SpRYn	NG NGN NNN		A₁-A₁₄			编辑效率高, 几近无PAM靶向, 窗口广	Tan et al., 2022
	SpRY-ABE8e	ecTadA:ecTadA^*		nCas9^D10A	NNN		A₃-A₁₀			几近无PAM靶向	Ren et al., 2021b
	ABE7.10^F148A	ecTadA^F148A: ecTadA^*F148A		nCas9^D10A	NGG		A₅-A₆			活性窗口缩小至1-2 nt	Zhou et al., 2019
CG碱基编辑(GBEs)	GBE	AID rAPOBEC1		nCas9^D10A	NGG		C₃-C₇	C:G>A:T/G:C		实现嘧啶与嘌呤间的颠换	Zhao et al., 2021
CG碱基编辑(GBEs)	CGBE	rAPOBEC1		nCas9^D10A	NGG		C₅-C₆	C:G>A:T/G:C		实现嘧啶与嘌呤间的颠换	Chen et al., 2021
双碱基编辑(A&CBE)	A&C-BEmax	hAID ecTadA:ecTadA^*		nCas9^D10A	NGG		C₂-C₁₇ A₄-A₇	C:G>T:A A:T>G:C		实现A、C共编辑	Zhang et al., 2020
	Target-ACE	PmCDA1 ecTadA:ecTadA^*			NGG		C₁-C₁₀ A₄-A₈				Sakata et al., 2019
	SPACE	PmCDA1 ecTadA^*			NGG		C₂-C₇ A₄-A₇				Grünewald et al., 2020
	ACBE	PmCDA1 ecTadA:ecTadA^*			NGG		C₁-C₇ A₄-A₆				Xie et al., 2020
	STEME	hAPOBEC3A ecTadA:ecTadA^*			NG NGD		C₁-C₁₇ A₄-A₈			在植物中实现A、 C共编辑	Li et al., 2020a
	pDuBE1	TadA8e			NGG		C₁-C₁₀ A₂-A₉			在植物中实现A、 C共编辑	Xu et al., 2021
编辑技术	编辑器		脱氨酶/逆转录酶	Cas蛋白	PAM (5′-3′)	活性窗口			编辑类型	特点	参考文献
先导编辑(PEs)	PE		M-MLV RT	nCas9^H840A	NGG	1-50			12种碱基替换插入(<15 bp) 缺失(<40 bp)	不受PAM的距离限制	Anzalone et al., 2019
	PPE		CaMV RT Retron RT							在植物中实现PE 应用	Lin et al., 2020
	pPE2		M-MLV RT							在植物中实现PE 应用	Xu et al., 2020a
	ePPE		M-MLV RT^{∆RNase H} M-MLV RT:NC			1-92			12种碱基替换插入(<40 bp) 缺失(<100 bp)	提高了在植物中的编辑效率	Zong et al., 2022
多重编辑	SWISSs		rAPOBEC1 ecTadA:ecTadA^*	nCas9^D10A	NG	C₃-C₁₆ A₄-A₇			C:G>T:A A:T>G:C Indels	具有A>G、C>T和Indels三重编辑功能	Li et al., 2020b
片段删除编辑	AFIDs		hAPOBEC3A hAPOBEC3Bctd	Cas9	NGG	C₁-C₁₄			多核苷酸删除	精准高效、可预测的多核苷酸缺失	Wang et al., 2020

物种		品种	靶基因	转化方法	编辑技术	脱氨酶/逆转录酶		编辑类型	编辑窗口	编辑效率(%)		参考文献
水稻(Oryza sativa)		中花11	CDC48 NRT1.1B-T1	农杆菌AGL1	CBEs	hAPOBEC3A		C:G>T:A	C₂-C₁₆	44.1- 82.9		Zong et al., 2018
		日本晴	CDC48	农杆菌AGL1		rAPOBEC1		C:G>T:A	C₃-C₈	43.48		Zong et al., 2017
		Kitaake	Pi-d2 FLS2	农杆菌EHA105		hAID^*Δ		C:G>T:A	C₃-C₇	30.8-57		Ren et al., 2018
		中花11	ACC-T1 ALS-T1 CDC48-T3 DEP1-T1/2 NRT1.1B-T1	农杆菌AGL1	ABEs	ecTadA: ecTadA^*		A:T>G:C	A₄-A₈	15.8- 59.1		Li et al., 2018b
		日本晴	SPL14/16/17/18 SLR1	农杆菌EHA105		ecTadA: ecTadA^*		A:T>G:C	A₅-A₁₄	12.5- 61.3		Hua et al., 2018
		Kitaake	SERK2 WRKY45	农杆菌EHA105		ecTadA: ecTadA^*		A:T>G:C	A₄-A₇	32.05- 62.26		Yan et al., 2018
		中花11	ACC	农杆菌AGL1	STEMEs	hAPOBEC3A ecTadA: ecTadA^*		C:G>T:A; A:T>G:C	C₁-C₁₇ A₄-A₈	3.84		Li et al., 2020a
		中花11	CDC48-T1 ALS-T2	农杆菌EHA105	PEs	M-MLV		G:C>T:A 插入(≤3 bp) 删除(≤6 bp)	N₁-N₆	2.6-21.8		Lin et al., 2020
		日本晴	ALS-1/2 ACC-1 DEP1	农杆菌EHA105	PEs	M-MLV		C:G>T:A A:T>G:C G:C>T:A G:C>C:G G:C>A:T T:A>A:T A:T>C:G	N₁₈-N₃₃	1.7-26		Xu et al., 2020b
物种	品种		靶基因	转化方法	编辑技术	脱氨酶/逆转录酶	编辑类型		编辑窗口	编辑效率(%)	参考文献
	中花11		ALS-T2 ACC-T2 BADH-indels	农杆菌AGL1	SWISS	hAPOBEC3A ecTadA: ecTadA^*	C:G>T:A A:T>G:C C:G>G:C 删除(≤45 bp)		C₆-C₇ A₄-A₇	7.3	Li et al., 2020b
	中花11		CDC48-T2 SPL14 SWEET14	农杆菌AGL1	AFID	hAPOBEC3A	删除(≤16 bp)		C₂-C₁₇	22.2- 55.8	Wang et al., 2020
小麦(Triticum aestivum)	Kenong 199		ALS MTL	基因枪	CBEs	hAPOBEC3A	C:G>T:A		C_-9-C₁₃	16.7- 22.5	Zong et al., 2018
	Bobwhite		LOX2-S1	基因枪	CBEs	rAPOBEC1	C:G>T:A		C₃-C₉	1.25	Zong et al., 2017
	Kenong 199		ALS	基因枪	CBEs	rAPOBEC1	C:G>T:A		C_-1-C₇	22-78	Zhang et al., 2019
	Kenong 199		DEP1 GW2	基因枪	ABEs	ecTadA: ecTadA^*	A:T>G:C		A₅-A₈	0.4-1.1	Li et al., 2018a
	Kenong 199		miR396 GASR6	基因枪	AFID	hAPOBEC3A	删除(≤35 bp)		C_-12-C₂₃	25-37.5	Wang et al., 2020
番茄(Solanum lycopersicum)	WVA106		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₇	34.7	Veillet et al., 2019
	WVA106		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₁-C₈	20.59	Veillet et al., 2020
	Micro- Tom		DELLA	农杆菌	CBEs	PmCDA1	C:G>T:A		C₁-C₃	50.5	Shimatani et al., 2017
马铃薯 (S. tuberosum)	Désirée		GBSS1 DMR6-1	PEG	CBEs	hAPOBEC3A	C:G>T:A		C₃-C₁₀	8-16.67	Veillet et al., 2020
	Désirée		GBSS-T6	PEG	CBEs	hAPOBEC3A	C:G>T:A		C₁-C₁₃	6.5	Jiang et al., 2022
	Désirée		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₁-C₈	25	Veillet et al., 2019
大豆 (Glycine max)	Jack		FT2a FT4	农杆菌	CBEs	rAPOBEC1	C:G>T:A		C₆-C₇	5.88- 18.2	Cai et al., 2020
棉花(Gossypium hirsutum)	Jin668		CLA PEBP	农杆菌GV3101	CBEs	rAPOBEC1	C:G>T:A		C₄-C₈	26.67- 57.78	Qin et al., 2020
玉米 (Zea mays)	Zong31		CENH3	农杆菌AGL1	CBEs	rAPOBEC1	C:G>T:A		C₃-C₈	10.1	Zong et al., 2017
玉米 (Zea mays)	ND73		ALS1/2	农杆菌LBA4404/ pVS1-VIR2	PEs	CmYLCV	C:G>T:A A:T>C:G T:A>G:C T:A>C:G C:G>A:T G:C>T:A G:C>A:T G:C>C:G		N₃-N₄₆	6.5-53.2	Jiang et al., 2020
油菜(Brassica napus)	J9712		ALS1	农杆菌	CBEs	rAPOBEC1	C:G>T:A		C₅-C₇	1.8	Wu et al., 2020
西瓜(Citrullus lanatus)	ZG94		ALS	农杆菌EHA105	CBEs	rAPOBEC1	C:G>T:A		C₇-C₈	23	Tian et al., 2018

物种		品种	靶基因	转化方法	编辑技术	脱氨酶/逆转录酶		编辑类型	编辑窗口	编辑效率(%)		参考文献
水稻(Oryza sativa)		中花11	CDC48 NRT1.1B-T1	农杆菌AGL1	CBEs	hAPOBEC3A		C:G>T:A	C₂-C₁₆	44.1- 82.9		Zong et al., 2018
		日本晴	CDC48	农杆菌AGL1		rAPOBEC1		C:G>T:A	C₃-C₈	43.48		Zong et al., 2017
		Kitaake	Pi-d2 FLS2	农杆菌EHA105		hAID^*Δ		C:G>T:A	C₃-C₇	30.8-57		Ren et al., 2018
		中花11	ACC-T1 ALS-T1 CDC48-T3 DEP1-T1/2 NRT1.1B-T1	农杆菌AGL1	ABEs	ecTadA: ecTadA^*		A:T>G:C	A₄-A₈	15.8- 59.1		Li et al., 2018b
		日本晴	SPL14/16/17/18 SLR1	农杆菌EHA105		ecTadA: ecTadA^*		A:T>G:C	A₅-A₁₄	12.5- 61.3		Hua et al., 2018
		Kitaake	SERK2 WRKY45	农杆菌EHA105		ecTadA: ecTadA^*		A:T>G:C	A₄-A₇	32.05- 62.26		Yan et al., 2018
		中花11	ACC	农杆菌AGL1	STEMEs	hAPOBEC3A ecTadA: ecTadA^*		C:G>T:A; A:T>G:C	C₁-C₁₇ A₄-A₈	3.84		Li et al., 2020a
		中花11	CDC48-T1 ALS-T2	农杆菌EHA105	PEs	M-MLV		G:C>T:A 插入(≤3 bp) 删除(≤6 bp)	N₁-N₆	2.6-21.8		Lin et al., 2020
		日本晴	ALS-1/2 ACC-1 DEP1	农杆菌EHA105	PEs	M-MLV		C:G>T:A A:T>G:C G:C>T:A G:C>C:G G:C>A:T T:A>A:T A:T>C:G	N₁₈-N₃₃	1.7-26		Xu et al., 2020b
物种	品种		靶基因	转化方法	编辑技术	脱氨酶/逆转录酶	编辑类型		编辑窗口	编辑效率(%)	参考文献
	中花11		ALS-T2 ACC-T2 BADH-indels	农杆菌AGL1	SWISS	hAPOBEC3A ecTadA: ecTadA^*	C:G>T:A A:T>G:C C:G>G:C 删除(≤45 bp)		C₆-C₇ A₄-A₇	7.3	Li et al., 2020b
	中花11		CDC48-T2 SPL14 SWEET14	农杆菌AGL1	AFID	hAPOBEC3A	删除(≤16 bp)		C₂-C₁₇	22.2- 55.8	Wang et al., 2020
小麦(Triticum aestivum)	Kenong 199		ALS MTL	基因枪	CBEs	hAPOBEC3A	C:G>T:A		C_-9-C₁₃	16.7- 22.5	Zong et al., 2018
	Bobwhite		LOX2-S1	基因枪	CBEs	rAPOBEC1	C:G>T:A		C₃-C₉	1.25	Zong et al., 2017
	Kenong 199		ALS	基因枪	CBEs	rAPOBEC1	C:G>T:A		C_-1-C₇	22-78	Zhang et al., 2019
	Kenong 199		DEP1 GW2	基因枪	ABEs	ecTadA: ecTadA^*	A:T>G:C		A₅-A₈	0.4-1.1	Li et al., 2018a
	Kenong 199		miR396 GASR6	基因枪	AFID	hAPOBEC3A	删除(≤35 bp)		C_-12-C₂₃	25-37.5	Wang et al., 2020
番茄(Solanum lycopersicum)	WVA106		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₇	34.7	Veillet et al., 2019
	WVA106		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₁-C₈	20.59	Veillet et al., 2020
	Micro- Tom		DELLA	农杆菌	CBEs	PmCDA1	C:G>T:A		C₁-C₃	50.5	Shimatani et al., 2017
马铃薯 (S. tuberosum)	Désirée		GBSS1 DMR6-1	PEG	CBEs	hAPOBEC3A	C:G>T:A		C₃-C₁₀	8-16.67	Veillet et al., 2020
	Désirée		GBSS-T6	PEG	CBEs	hAPOBEC3A	C:G>T:A		C₁-C₁₃	6.5	Jiang et al., 2022
	Désirée		ALS1	农杆菌C58 pGV2260	CBEs	PmCDA1	C:G>T:A		C₁-C₈	25	Veillet et al., 2019
大豆 (Glycine max)	Jack		FT2a FT4	农杆菌	CBEs	rAPOBEC1	C:G>T:A		C₆-C₇	5.88- 18.2	Cai et al., 2020
棉花(Gossypium hirsutum)	Jin668		CLA PEBP	农杆菌GV3101	CBEs	rAPOBEC1	C:G>T:A		C₄-C₈	26.67- 57.78	Qin et al., 2020
玉米 (Zea mays)	Zong31		CENH3	农杆菌AGL1	CBEs	rAPOBEC1	C:G>T:A		C₃-C₈	10.1	Zong et al., 2017
玉米 (Zea mays)	ND73		ALS1/2	农杆菌LBA4404/ pVS1-VIR2	PEs	CmYLCV	C:G>T:A A:T>C:G T:A>G:C T:A>C:G C:G>A:T G:C>T:A G:C>A:T G:C>C:G		N₃-N₄₆	6.5-53.2	Jiang et al., 2020
油菜(Brassica napus)	J9712		ALS1	农杆菌	CBEs	rAPOBEC1	C:G>T:A		C₅-C₇	1.8	Wu et al., 2020
西瓜(Citrullus lanatus)	ZG94		ALS	农杆菌EHA105	CBEs	rAPOBEC1	C:G>T:A		C₇-C₈	23	Tian et al., 2018