CRISPR/Cas9技术研究进展及其在园艺植物中的应用进展

doi:10.16420/j.issn.0513-353x.2023-0760

摘要/Abstract

摘要：

CRISPR/Cas9是细菌和古细菌的免疫防御系统，1987年首次在大肠杆菌中发现一段特殊的重复间隔序列，后来在20多种细菌和古细菌中发现此重复间隔序列，2002年这段特殊的序列被正式命名为CRISPR，之后人们便展开了一系列利用CRISPR/Cas9技术进行基因编辑的研究。CRISPR/Cas9技术是继锌指核酸酶（zinc finger nucleases，ZFNs）技术、转录激活因子样效应核酸酶（transcription on activator-like effector nucleases，TALENs）技术后第三代基因编辑技术。该系统具有操作设计简单，突变效率高，成本低等优点，陆续应用在柑橘、葡萄、香蕉、草莓、黄瓜和马铃薯等众多园艺植物中。本文综述了CRISPR/Cas9技术的作用原理及其研究进展，论述了各种编辑器的发展历程，包括单碱基编辑器、双碱基编辑器及引导编辑器，介绍了CRISPR/Cas9技术在园艺植物中的应用，提出其存在的问题以及未来展望。

关键词: CRISPR/Cas9, 基因编辑器, 研究进展, 园艺植物

Abstract:

CRISPR/Cas9 is the immune defense system in bacteria and archaea. In 1987，a special repeat interval sequence was firstly discovered in Escherichia coli. Later，this repeat interval sequence was also found in more than 20 bacteria and archaea. In 2002，this special sequence was officially named as CRISPR. Subsequently，a series of studies using CRISPR/Cas9 technology for gene editing were carried out. CRISPR/Cas9 technology is the third generation gene editing technology，following ZFNs（zinc finger nucleases）technology and TALENs（transfer activator like effector nucleases）technology. This system has the advantages of simple operation design，high mutation efficiency，low cost，and has been successively applied in many horticultural plants such as citrus，grape，banana，strawberry，cucumber，and potato. This article reviews the principles and research progress of CRISPR/Cas9 technology，discusses the developmental history of various editors，including single base editor，double base editor，and guided editor，introduces the application of CRISPR/Cas9 technology in horticultural plants，and finally proposes the remaining problems and future prospects of CRISPR/Cas9 technology.

Key words: CRISPR/Cas9, gene editor, research progress, horticultural plants

王晨雨, 刘孟军, 王立新, 刘志国. CRISPR/Cas9技术研究进展及其在园艺植物中的应用进展[J]. 园艺学报, 2024, 51(7): 1439-1454.

WANG Chenyu, LIU Mengjun, WANG Lixin, LIU Zhiguo. Research Progress of CRISPR/Cas9 Technology and Its Application in Horticultural Plants[J]. Acta Horticulturae Sinica, 2024, 51(7): 1439-1454.

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应用 Application	种类 Species	目标基因 Target gene	目标性状 Target trait	参考文献 Reference
调节生长特性 Regulating growth characteristics	苹果 Malus × domestica	MdPDS	白化表型 Albino phenotype	Charrier et al.，2019
	香蕉 Musa nana Lour.	MaGA20ox2	半矮生 Semidwarf	Shao et al.，2020
	西瓜 Citrullus lanatus	ClBG1	改变种子大小和质量Changing seed size and quality	Wang et al.，2021
	蓝莓 Vaccinium spp.	CEN	促营养生长Promote nutritional growth	Omori et al.，2021
	番茄 Solanum lycopersicum	AGL6	单性结实 Parthenocarpy	Klap et al.，2017
		MBP21	果柄无分离层Fruit stalk without separation layer	Roldan et al.，2017
		SlTM6	雄性不育系 male sterile line	杨亮等，2023
	黄瓜 Cucumis sativus	CsVFB1	矮生 Dwarf	戚晶晶，2017
		CsCLAVATA3	改变心皮数Change the number of carpels	秦楠楠，2020
		Cs SH1	短下胚轴 Short hypocotyl	伍若彤，2021
	百合Lilium	LpPDS	白化表型 Albino phenotype	Yan et al.，2019
	菊花 Chrysanthemum	AN4	株形、叶片面积、冠径、株高和色素含量 Plant shape，leaf area，crown diameter，plant height，and pigment content	张翠，2021
改变开花时间及花色 Change flowering time and color	柑橘 Citrus reticulata Blanco	DUO1	改变花粉育性 Changing pollen fertility	徐彦辉，2023
	番茄 Solanum lycopersicum	SlFAF	调节开花 Adjusting flowering	Shang et al.，2024
	番茄 Solanum lycopersicum	TMF	花序简化 Simplified inflorescence	Xu et al.，2016
	日本牵牛花 Ipomoea nil	DFR-B	获得白色花 Obtain white flowers	Watanabe et al.，2017
		CCD4	白色花瓣变成淡黄色 White petals turn pale yellow	Watanabe et al.，2018
		Gt5GT，Gt3’GT，Gt5/3’AT	获得淡红色、紫罗兰色、暗粉红色和淡紫色花 Get light red，violet，dark pink，and light purple flowers	Tasaki et al.，2019
		GST1	获得白色和淡蓝色花Obtain white and light blue flowers	Tasaki et al.，2020
	矮牵牛 Petunia hybrid Vilm.	ACO1	延长花寿命Extend the lifespan of flowers	Xu et al.，2020
		AN4	花冠管脉络缺失Lacking veins in the corolla tube	Zhang et al.，2021
		miRl59b	延迟开花 Delayed flowering	王浩，2019
	蓝猪耳 Torenia fournieri	TfRAD1	改变花瓣形状和颜色Change petal shape and color	Su et al.，2017
	蓝猪耳 Torenia fournieri	F3H	花瓣改色 Petal color change	Nishihara et al.，2018
提高果实品质 Improve fruit quality	香蕉 Musa nana Lour.	LCYε	提高β胡萝卜素Increase β carotene	Kaur et al.，2020
	草莓 Fragaria × ananassa	FaPG1	提高硬度 Improve hardness	López-Casado et al.，2023
		FvCO4	果实发育密切相关Fruit development is closely related	陈文君，2019
		PHO2	提高果实中磷含量 Increase the phosphorus content in fruits	Zhang et al.，2023
		RAP	绿茎白果 Green stemmed white fruit	Gao et al.，2020
	毛酸浆 Physalis pubescens	SELF-PRUNING 5G	提高果实数量和质量 Improve the quantity and quality of fruits	Lemmon et al.，2018
	番茄 Solanum lycopersicum	CRITOSO	橙色果实 Orange fruit	Isaacson et al.，2002
		MYB12	粉红色果实 Pink fruit	Ballester et al.，2010
		PSY1	黄色果实 Yellow fruit	Filler et al.，2017
		RIN	果实成熟延迟 Delayed fruit ripening	Lang et al.，2017
		SlACS2	果实成熟延迟 Delayed fruit ripening	刘江娜等，2023
		SlbZIP1	提高糖和氨基酸含量 Increase sugar and amino acid content	Nguyen et al.，2023
		SNAC9	果实成熟延迟 Delayed fruit ripening	Feng et al.，2023
增强植株抗性 Enhance plant resistance	苹果 Malus × domestica	FB_MR5	火枯病症状下降Reduced symptoms of fire blight	Broggini et al.，2014
		FB_MR5	增强抗火枯病能力Enhance the ability to resist fire blight	Kost et al.，2015
		HIPM	火疫病减轻Alleviation of fire epidemic	Campa et al.，2019
	柑橘 Citrus reticulata Blanco	CcPUB4	增强耐盐性Enhance salt resistance	李秋月，2019
	柑橘 Citrus reticulata Blanco	CsLOB1	抗柑橘溃疡病Anti citrus ulcer disease	Hu et al.，2014
	葡萄 Vitis vinifera	VvBAK1，VvLecRK1	抗白粉病 Anti powdery mildew	贾慧，2019
		VvWRKY52	抗灰葡萄孢Resistance to botrytis cinerea	Wang et al.，2018
		VvPR4b	抗霜霉病Anti downy mildew disease	李梦媛，2020
		VviEDR2	抗白粉病 Anti powdery mildew	杨禄山等，2020
增强植株抗性 Enhance plant resistance	香蕉Musa nana Lour.	eBSV	抗条斑病毒 Anti stripe virus	Tripathi et al.，2019
	香蕉Musa nana Lour.	MusaDMR6	抗黄萎病 Verticillium wilt	Tripathi et al.，2021
	西瓜 Citrullus lanatus	ALS	抗除草剂 Herbicide resistant	张倩倩，2019
	西瓜 Citrullus lanatus	Clpsk1	抗尖孢镰刀菌Anti Fusarium oxysporum	Zhang et al.，2020a
	草莓 Fragaria × ananassa	FVICE1	抗严寒和干旱Resistance to severe cold and drought	Han et al.，2023
	番茄 Solanum lycopersicum	BZR1	参与调控耐热途径 Participate in regulating heat resistance pathways	Yin et al.，2018
		Mlo1	抗白粉病 Anti powdery mildew	Nekrasov et al.，2017
		MAPK3	抗灰霉病、耐旱Resistant to gray mold and drought	Zhang et al.，2018；Wang et al.，2017
		SlCBF1	防止植物受冻害Prevent plants from freezing damage	Li et al.，2018
		SINPR1	耐旱 Drought resistance	Li et al.，2019
	黄瓜 Cucumis sativus	eIF4E	抗黄瓜叶脉黄变病毒、西葫芦黄花叶病毒、番木瓜环斑花叶病毒Anti cucumber vein yellowing virus，zucchini yellow mosaic virus，papaya ring spot mosaic virus	Chandrasekaran et al.，2016
	黄瓜 Cucumis sativus	Mlo1	抗白粉病 Anti powdery mildew	Shnaider et al.，2023
	马铃薯 Solanum tuberosum	PVY	抗马铃薯Y病毒 Anti potato Y Virus	Lucioli et al.，2022
		SmelPPO	降低褐化 Reduce browning	González et al.，2020
		StCHLI	抗晚疫病 Anti late epidemic disease	Kieu et al.，2021
		St16DOX	减少糖苷生物碱Reduce glycoside alkaloids	Nakayasu et al.，2018
	茄子 Solanum melongena	SmelPPO	降低褐化 Reduce browning	Maioli et al.，2020

应用 Application	种类 Species	目标基因 Target gene	目标性状 Target trait	参考文献 Reference
调节生长特性 Regulating growth characteristics	苹果 Malus × domestica	MdPDS	白化表型 Albino phenotype	Charrier et al.，2019
	香蕉 Musa nana Lour.	MaGA20ox2	半矮生 Semidwarf	Shao et al.，2020
	西瓜 Citrullus lanatus	ClBG1	改变种子大小和质量Changing seed size and quality	Wang et al.，2021
	蓝莓 Vaccinium spp.	CEN	促营养生长Promote nutritional growth	Omori et al.，2021
	番茄 Solanum lycopersicum	AGL6	单性结实 Parthenocarpy	Klap et al.，2017
		MBP21	果柄无分离层Fruit stalk without separation layer	Roldan et al.，2017
		SlTM6	雄性不育系 male sterile line	杨亮等，2023
	黄瓜 Cucumis sativus	CsVFB1	矮生 Dwarf	戚晶晶，2017
		CsCLAVATA3	改变心皮数Change the number of carpels	秦楠楠，2020
		Cs SH1	短下胚轴 Short hypocotyl	伍若彤，2021
	百合Lilium	LpPDS	白化表型 Albino phenotype	Yan et al.，2019
	菊花 Chrysanthemum	AN4	株形、叶片面积、冠径、株高和色素含量 Plant shape，leaf area，crown diameter，plant height，and pigment content	张翠，2021
改变开花时间及花色 Change flowering time and color	柑橘 Citrus reticulata Blanco	DUO1	改变花粉育性 Changing pollen fertility	徐彦辉，2023
	番茄 Solanum lycopersicum	SlFAF	调节开花 Adjusting flowering	Shang et al.，2024
	番茄 Solanum lycopersicum	TMF	花序简化 Simplified inflorescence	Xu et al.，2016
	日本牵牛花 Ipomoea nil	DFR-B	获得白色花 Obtain white flowers	Watanabe et al.，2017
		CCD4	白色花瓣变成淡黄色 White petals turn pale yellow	Watanabe et al.，2018
		Gt5GT，Gt3’GT，Gt5/3’AT	获得淡红色、紫罗兰色、暗粉红色和淡紫色花 Get light red，violet，dark pink，and light purple flowers	Tasaki et al.，2019
		GST1	获得白色和淡蓝色花Obtain white and light blue flowers	Tasaki et al.，2020
	矮牵牛 Petunia hybrid Vilm.	ACO1	延长花寿命Extend the lifespan of flowers	Xu et al.，2020
		AN4	花冠管脉络缺失Lacking veins in the corolla tube	Zhang et al.，2021
		miRl59b	延迟开花 Delayed flowering	王浩，2019
	蓝猪耳 Torenia fournieri	TfRAD1	改变花瓣形状和颜色Change petal shape and color	Su et al.，2017
	蓝猪耳 Torenia fournieri	F3H	花瓣改色 Petal color change	Nishihara et al.，2018
提高果实品质 Improve fruit quality	香蕉 Musa nana Lour.	LCYε	提高β胡萝卜素Increase β carotene	Kaur et al.，2020
	草莓 Fragaria × ananassa	FaPG1	提高硬度 Improve hardness	López-Casado et al.，2023
		FvCO4	果实发育密切相关Fruit development is closely related	陈文君，2019
		PHO2	提高果实中磷含量 Increase the phosphorus content in fruits	Zhang et al.，2023
		RAP	绿茎白果 Green stemmed white fruit	Gao et al.，2020
	毛酸浆 Physalis pubescens	SELF-PRUNING 5G	提高果实数量和质量 Improve the quantity and quality of fruits	Lemmon et al.，2018
	番茄 Solanum lycopersicum	CRITOSO	橙色果实 Orange fruit	Isaacson et al.，2002
		MYB12	粉红色果实 Pink fruit	Ballester et al.，2010
		PSY1	黄色果实 Yellow fruit	Filler et al.，2017
		RIN	果实成熟延迟 Delayed fruit ripening	Lang et al.，2017
		SlACS2	果实成熟延迟 Delayed fruit ripening	刘江娜等，2023
		SlbZIP1	提高糖和氨基酸含量 Increase sugar and amino acid content	Nguyen et al.，2023
		SNAC9	果实成熟延迟 Delayed fruit ripening	Feng et al.，2023
增强植株抗性 Enhance plant resistance	苹果 Malus × domestica	FB_MR5	火枯病症状下降Reduced symptoms of fire blight	Broggini et al.，2014
		FB_MR5	增强抗火枯病能力Enhance the ability to resist fire blight	Kost et al.，2015
		HIPM	火疫病减轻Alleviation of fire epidemic	Campa et al.，2019
	柑橘 Citrus reticulata Blanco	CcPUB4	增强耐盐性Enhance salt resistance	李秋月，2019
	柑橘 Citrus reticulata Blanco	CsLOB1	抗柑橘溃疡病Anti citrus ulcer disease	Hu et al.，2014
	葡萄 Vitis vinifera	VvBAK1，VvLecRK1	抗白粉病 Anti powdery mildew	贾慧，2019
		VvWRKY52	抗灰葡萄孢Resistance to botrytis cinerea	Wang et al.，2018
		VvPR4b	抗霜霉病Anti downy mildew disease	李梦媛，2020
		VviEDR2	抗白粉病 Anti powdery mildew	杨禄山等，2020
增强植株抗性 Enhance plant resistance	香蕉Musa nana Lour.	eBSV	抗条斑病毒 Anti stripe virus	Tripathi et al.，2019
	香蕉Musa nana Lour.	MusaDMR6	抗黄萎病 Verticillium wilt	Tripathi et al.，2021
	西瓜 Citrullus lanatus	ALS	抗除草剂 Herbicide resistant	张倩倩，2019
	西瓜 Citrullus lanatus	Clpsk1	抗尖孢镰刀菌Anti Fusarium oxysporum	Zhang et al.，2020a
	草莓 Fragaria × ananassa	FVICE1	抗严寒和干旱Resistance to severe cold and drought	Han et al.，2023
	番茄 Solanum lycopersicum	BZR1	参与调控耐热途径 Participate in regulating heat resistance pathways	Yin et al.，2018
		Mlo1	抗白粉病 Anti powdery mildew	Nekrasov et al.，2017
		MAPK3	抗灰霉病、耐旱Resistant to gray mold and drought	Zhang et al.，2018；Wang et al.，2017
		SlCBF1	防止植物受冻害Prevent plants from freezing damage	Li et al.，2018
		SINPR1	耐旱 Drought resistance	Li et al.，2019
	黄瓜 Cucumis sativus	eIF4E	抗黄瓜叶脉黄变病毒、西葫芦黄花叶病毒、番木瓜环斑花叶病毒Anti cucumber vein yellowing virus，zucchini yellow mosaic virus，papaya ring spot mosaic virus	Chandrasekaran et al.，2016
	黄瓜 Cucumis sativus	Mlo1	抗白粉病 Anti powdery mildew	Shnaider et al.，2023
	马铃薯 Solanum tuberosum	PVY	抗马铃薯Y病毒 Anti potato Y Virus	Lucioli et al.，2022
		SmelPPO	降低褐化 Reduce browning	González et al.，2020
		StCHLI	抗晚疫病 Anti late epidemic disease	Kieu et al.，2021
		St16DOX	减少糖苷生物碱Reduce glycoside alkaloids	Nakayasu et al.，2018
	茄子 Solanum melongena	SmelPPO	降低褐化 Reduce browning	Maioli et al.，2020

种类 Species	碱基编辑器 Base editor	目标基因 Target gene	目标性状 Target trait	编辑效率/% Editing efficiency	参考文献 Reference
西瓜 Citrullus lanatus	BE3	ClALS	抗除草剂 Herbicide resistant	23.0	Tian et al.，2018
番茄 Solanum lycopersicum	Target-AID	DELLA，ETR1	株型 Plant type	50.5	Shimatani et al.，2017
	CBE	ALS	抗除草剂 Herbicide resistant	71.0	Veillet et al.，2019b
	pCXPE03	GAI，ALS2，PDS1	抗除草剂 Herbicide resistant	3.4 ~ 6.7	Lu et al.，2021
马铃薯 Solanum tuberosum	A3A-PBE	StGBSS-T6	淀粉含量 Starch content	6.5	Zong et al.，2018
马铃薯 Solanum tuberosum	CBE	StGBSS1	淀粉含量 Starch content	8.00 ~ 16.67	Veillet et al.，2019a
油菜 Brassica napus	CBE	BnALS1，BnALS3	抗除草剂 Herbicide resistant	1.8	Wu et al.，2020

种类 Species	碱基编辑器 Base editor	目标基因 Target gene	目标性状 Target trait	编辑效率/% Editing efficiency	参考文献 Reference
西瓜 Citrullus lanatus	BE3	ClALS	抗除草剂 Herbicide resistant	23.0	Tian et al.，2018
番茄 Solanum lycopersicum	Target-AID	DELLA，ETR1	株型 Plant type	50.5	Shimatani et al.，2017
	CBE	ALS	抗除草剂 Herbicide resistant	71.0	Veillet et al.，2019b
	pCXPE03	GAI，ALS2，PDS1	抗除草剂 Herbicide resistant	3.4 ~ 6.7	Lu et al.，2021
马铃薯 Solanum tuberosum	A3A-PBE	StGBSS-T6	淀粉含量 Starch content	6.5	Zong et al.，2018
马铃薯 Solanum tuberosum	CBE	StGBSS1	淀粉含量 Starch content	8.00 ~ 16.67	Veillet et al.，2019a
油菜 Brassica napus	CBE	BnALS1，BnALS3	抗除草剂 Herbicide resistant	1.8	Wu et al.，2020