植物启动子研究进展

doi:10.16420/j.issn.0513-353x.2024-0408

摘要/Abstract

摘要：

探讨了不同类型启动子的结构特性及其在基因功能研究、代谢工程和基因编辑（CRISPR/ Cas9）中的作用，并总结了启动子研究的内容及方法，以期为深入理解和研究植物启动子的结构特性和调控机制提供帮助。

Abstract:

In this paper， the structural characteristics of different types of promoters and their roles in gene function research，metabolic engineering，and gene editing（CRISPR/Cas9）were discusses，and summarized the content and methods for promoter identification，aiming to provide insights for a deeper understanding and investigation of the structural characteristics and regulatory mechanisms of plant promoters.

Key words: promoter, genetic engineering, metabolic engineering, CRISPR/Cas9

张志愿, 王丹, 惠竹梅, 王现行. 植物启动子研究进展[J]. 园艺学报, 2025, 52(4): 1065-1092.

ZHANG Zhiyuan, WANG Dan, XI Zhumei, WANG Xianhang. Advances in Plant Promoter Research[J]. Acta Horticulturae Sinica, 2025, 52(4): 1065-1092.

图/表 7

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启动子类型 Promoter type	来源基因 Source gene	顺式作用元件 cis-Acting element	来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
根特异性启动子 Root-specific promoter	OsGT1	ROOTMOTIFTAPOX1、POLLEN1LELAT52	水稻 Oryza sativa	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2007b
	GhRACK1	ROOTMOTIFTAPOX1、MYB2、TBOX、LIBOX、IBOXCORE	陆地棉Gossypium hirsutum	烟草Nicotiana tabacum	GUS	杨江涛等,2016
	DcMYB113	/	胡萝卜 Daucus carota subsp. sativus	胡萝卜 Daucus carota subsp. sativus	LUC	Xu et al.,2020
	At1g	/	拟南芥 Arabidopsis thaliana	烟草Nicotiana tabacum	GUS	Ly et al.,2022
	rRSP1	ROOTMOTIFTAPOX1、CACTFTPPCA1	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Huang et al.,2015
	rRSP3	ROOTMOTIFTAPOX1、CACTFTPPCA1	水稻 Oryza sativa	水稻 Oryza sativa	GUS
	rRSP5	ROOTMOTIFTAPOX1、CACTFTPPCA1、OSE1/2ROOTNODULE	水稻 Oryza sativa	水稻 Oryza sativa	GUS
	SynR2	ATATT、MAR-box	人工合成 Synthetic	烟草 Nicotiana tabacum	GUS	Mohan et al.,2017
	E543254P	RHERPATEXPA7、SP8BFIBSP8BIB、OSE1ROOTNODULE、WUSATAg	番茄 Solanum lycopersicum	拟南芥 Arabidopsis thaliana	GUS	Lim et al.,2012
	E542814P	RHERPATEXPA7、SP8BFIBSP8BIB、OSE1ROOTNODULE、WUSATAg	番茄 Solanum lycopersicum	拟南芥 Arabidopsis thaliana	GUS	Lim et al.,2012
	GmTIP	OSE2ROOTNODULE、ROOTMOTIFTAPOX1	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Chen et al.,2015
	GmADR1	M1-motif、M2-motif、M3-motif、RHE	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Xun et al.,2021
	GmBTP1	M1-motif、M2-motif 、M3-motif、RHE	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS
	pNtREL1	OSE1、OSE2、ROOTMOTIFTAPOX1、SURECOREATSULTR11、P1BS、WUSATAg	烟草 Nicotiana tabacum	烟草 Nicotiana tabacum	GUS	Zhang et al.,2016
	Pyk10	ocs-elements、mas-elements、as-1-elements	拟南芥 Arabidopsis thaliana	拟南芥Arabidopsis thaliana	GUS	Nitz et al.,2001
花器官特异启动子 Floral organ-specific promoter	PsEND1	/	豌豆Pisum sativum	拟南芥Arabidopsis thaliana	GUS	Roque et al.,2019
	BraA08g014780.3C	POLLEN1LELAT52、GTGANTG10	白菜Brassica rapa	拟南芥Arabidopsis thaliana	GUS	Zhao et al.,2022
	CpMDAR4	POLLEN1LELAT52、CREs	木瓜 Carica papaya	拟南芥 Arabidopsis thaliana	GUS	Zerpa-Catanho et al.,2022
	rMdAG2I	TATA-box	人工合成 Synthetic	烟草、苹果 Nicotiana tabacum、Malus domestica	GUS	Dong et al.,2019
	JcTM6	CArG-box、GTGA、AGAAA-motif	麻风树 Jatropha curcas	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2020a
	PSG076	/	小麦 Triticum aestivum	小麦 Triticum aestivum	GUS	Chen et al.,2023b
	OsLSP3	LAT enhancer element、TGGTTA-motif、AGAAA-motif、AGGTCA-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP	Wang et al.,2020b
	OsLSP5	ATATT-motif、TTTCT-motif、AGAAA-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP
	OsLSP6	TGTGG-motif、TGGTTA-motif、AGAAA-motif、TTTCT-motif、AGGTCA-motif、AAATGA-motif、ATATT-motif、GATAA-motif、TAAATC-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP
	CtFSG2	ARE-box、Box4、CGTCA-motif、LTR	红花 Carthamus tinctorius	拟南芥 Arabidopsis thaliana	GUS	Chen et al.,2023a
种子特异性启动子 Seed-specific promoter	AtBGLU19	RY-motif、SEF3、SEF4、ESP、2SSEEDPROTBANAPA	拟南芥 Arabidopsis thaliana	拟南芥、烟草 Arabidopsis thaliana、Nicotiana tabacum	GUS	Liu et al.,2021c
	AX795651	/	陆地棉 Gossypium hirsutum	陆地棉 Gossypium hirsutum	GhDGAT1
	Emb5	AGCCCA-motif、GAAAAA-motif、CACT-motif、GCAA-motif、ABRE、TCA、Skn-1-motif、G-box、E-box、P-box、RAA、ATTTTC-motif、TGAATG-motif	玉米 Zea mays	拟南芥Arabidopsis thaliana	GUS	Li et al.,2021
	AHSSP2	RY-motif、G-box、GT1-box、EBOXBNNAPA、SEF1、SEF4	花生 Arachis hypogaea	拟南芥Arabidopsis thaliana	GUS	Yuan et al.,2019
	VvβVPE	CATGCA-motif、CAAACAC-motif、E-box、G-box	葡萄 Vitis vinifera	拟南芥Arabidopsis thaliana	GUS	Gong et al.,2019
果实特异性启动子 Fruit-specific promoter	2A11	F-1、F-2、F-3、F-4、F-5、S-1、S-2、S-3	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	van Haaren & Houck,1991,1993
	E4、E8	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	LUC、GUS	Kneissl & Deikman,1996；Xu et al.,1996
	PG	/	番茄、苹果 Solanum lycopersicum、Malus × domestica	番茄 Solanum lycopersicum	GUS	Montgomery et al.,1993b
	SlPR10	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GFP-GUS	Ruan et al.,2024
	SlAdh2	MBs、Box4、ARE	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Chung et al.,2020
	SlHDC-A	SBP-box、ATTTA-motif、GGATG-motif	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Kim et al.,2019
	pSolyc03g098700.1、pSolyc04g007770.2、pSolyc07g049140.2、pSolyc10g075090.1	(CA)n、ERE、GASRE、ARE2	番茄 Solanum lycopersicum	/	/	Chen et al.,2021b
	CitSEP	ARE、HSE、Box4、GCN4 motif、I-box	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS	Dasgupta et al.,2020
	CitWAX	HSE、Skn-1 motif、I-box、TCA element	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	CitJuSac	Box4、HSE、ABRE、Box1	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	CitVO1	GARE-motif、Box1、HSE、ARE、ABER	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	PamMybAp	ARE、HSE、BOx4、as-2-box、MBS	李子 Prunus salicina	番茄 Solanum lycopersicum	GUS
	GSX7R	ACE、Box4、CAT box、G-Box、G-box、GATA motif、GC motif、I-box、Sp1、chs-unit1 M1	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Lin et al.,2022
	AraSSU	/	拟南芥 Arabidopsis thaliana	鹰嘴豆 Cicer arietinum	cry2Aa	Boruah et al.,2023
	BiGSSP2	POsrbcs-62、OsAct1、POsrbcs-550	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS	Bai et al.,2020
	BiGSSP3	POsrbcs-62、OsAct1、4×GEAT	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
	BiGSSP7	POsrbcs-550、PD540-544、OsAct1、OsTub6I	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
	BiGSSP6	OsAct1	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
维管束特异性启动子 Vascular tissue-specific promoter	DROT1	/	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Sun et al.,2022
	SUC2	/	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Archer et al.,2023
	GmTIP1;6	MBS、ABRE、CGTCA-motif、TGACG-motif、TATC-box、TGA	毛豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Feng et al.,2022

启动子类型 Promoter type	来源基因 Source gene	顺式作用元件 cis-Acting element	来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
根特异性启动子 Root-specific promoter	OsGT1	ROOTMOTIFTAPOX1、POLLEN1LELAT52	水稻 Oryza sativa	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2007b
	GhRACK1	ROOTMOTIFTAPOX1、MYB2、TBOX、LIBOX、IBOXCORE	陆地棉Gossypium hirsutum	烟草Nicotiana tabacum	GUS	杨江涛等,2016
	DcMYB113	/	胡萝卜 Daucus carota subsp. sativus	胡萝卜 Daucus carota subsp. sativus	LUC	Xu et al.,2020
	At1g	/	拟南芥 Arabidopsis thaliana	烟草Nicotiana tabacum	GUS	Ly et al.,2022
	rRSP1	ROOTMOTIFTAPOX1、CACTFTPPCA1	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Huang et al.,2015
	rRSP3	ROOTMOTIFTAPOX1、CACTFTPPCA1	水稻 Oryza sativa	水稻 Oryza sativa	GUS
	rRSP5	ROOTMOTIFTAPOX1、CACTFTPPCA1、OSE1/2ROOTNODULE	水稻 Oryza sativa	水稻 Oryza sativa	GUS
	SynR2	ATATT、MAR-box	人工合成 Synthetic	烟草 Nicotiana tabacum	GUS	Mohan et al.,2017
	E543254P	RHERPATEXPA7、SP8BFIBSP8BIB、OSE1ROOTNODULE、WUSATAg	番茄 Solanum lycopersicum	拟南芥 Arabidopsis thaliana	GUS	Lim et al.,2012
	E542814P	RHERPATEXPA7、SP8BFIBSP8BIB、OSE1ROOTNODULE、WUSATAg	番茄 Solanum lycopersicum	拟南芥 Arabidopsis thaliana	GUS	Lim et al.,2012
	GmTIP	OSE2ROOTNODULE、ROOTMOTIFTAPOX1	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Chen et al.,2015
	GmADR1	M1-motif、M2-motif、M3-motif、RHE	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Xun et al.,2021
	GmBTP1	M1-motif、M2-motif 、M3-motif、RHE	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS
	pNtREL1	OSE1、OSE2、ROOTMOTIFTAPOX1、SURECOREATSULTR11、P1BS、WUSATAg	烟草 Nicotiana tabacum	烟草 Nicotiana tabacum	GUS	Zhang et al.,2016
	Pyk10	ocs-elements、mas-elements、as-1-elements	拟南芥 Arabidopsis thaliana	拟南芥Arabidopsis thaliana	GUS	Nitz et al.,2001
花器官特异启动子 Floral organ-specific promoter	PsEND1	/	豌豆Pisum sativum	拟南芥Arabidopsis thaliana	GUS	Roque et al.,2019
	BraA08g014780.3C	POLLEN1LELAT52、GTGANTG10	白菜Brassica rapa	拟南芥Arabidopsis thaliana	GUS	Zhao et al.,2022
	CpMDAR4	POLLEN1LELAT52、CREs	木瓜 Carica papaya	拟南芥 Arabidopsis thaliana	GUS	Zerpa-Catanho et al.,2022
	rMdAG2I	TATA-box	人工合成 Synthetic	烟草、苹果 Nicotiana tabacum、Malus domestica	GUS	Dong et al.,2019
	JcTM6	CArG-box、GTGA、AGAAA-motif	麻风树 Jatropha curcas	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2020a
	PSG076	/	小麦 Triticum aestivum	小麦 Triticum aestivum	GUS	Chen et al.,2023b
	OsLSP3	LAT enhancer element、TGGTTA-motif、AGAAA-motif、AGGTCA-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP	Wang et al.,2020b
	OsLSP5	ATATT-motif、TTTCT-motif、AGAAA-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP
	OsLSP6	TGTGG-motif、TGGTTA-motif、AGAAA-motif、TTTCT-motif、AGGTCA-motif、AAATGA-motif、ATATT-motif、GATAA-motif、TAAATC-motif	水稻 Oryza sativa	水稻 Oryza sativa	GFP
	CtFSG2	ARE-box、Box4、CGTCA-motif、LTR	红花 Carthamus tinctorius	拟南芥 Arabidopsis thaliana	GUS	Chen et al.,2023a
种子特异性启动子 Seed-specific promoter	AtBGLU19	RY-motif、SEF3、SEF4、ESP、2SSEEDPROTBANAPA	拟南芥 Arabidopsis thaliana	拟南芥、烟草 Arabidopsis thaliana、Nicotiana tabacum	GUS	Liu et al.,2021c
	AX795651	/	陆地棉 Gossypium hirsutum	陆地棉 Gossypium hirsutum	GhDGAT1
	Emb5	AGCCCA-motif、GAAAAA-motif、CACT-motif、GCAA-motif、ABRE、TCA、Skn-1-motif、G-box、E-box、P-box、RAA、ATTTTC-motif、TGAATG-motif	玉米 Zea mays	拟南芥Arabidopsis thaliana	GUS	Li et al.,2021
	AHSSP2	RY-motif、G-box、GT1-box、EBOXBNNAPA、SEF1、SEF4	花生 Arachis hypogaea	拟南芥Arabidopsis thaliana	GUS	Yuan et al.,2019
	VvβVPE	CATGCA-motif、CAAACAC-motif、E-box、G-box	葡萄 Vitis vinifera	拟南芥Arabidopsis thaliana	GUS	Gong et al.,2019
果实特异性启动子 Fruit-specific promoter	2A11	F-1、F-2、F-3、F-4、F-5、S-1、S-2、S-3	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	van Haaren & Houck,1991,1993
	E4、E8	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	LUC、GUS	Kneissl & Deikman,1996；Xu et al.,1996
	PG	/	番茄、苹果 Solanum lycopersicum、Malus × domestica	番茄 Solanum lycopersicum	GUS	Montgomery et al.,1993b
	SlPR10	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GFP-GUS	Ruan et al.,2024
	SlAdh2	MBs、Box4、ARE	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Chung et al.,2020
	SlHDC-A	SBP-box、ATTTA-motif、GGATG-motif	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Kim et al.,2019
	pSolyc03g098700.1、pSolyc04g007770.2、pSolyc07g049140.2、pSolyc10g075090.1	(CA)n、ERE、GASRE、ARE2	番茄 Solanum lycopersicum	/	/	Chen et al.,2021b
	CitSEP	ARE、HSE、Box4、GCN4 motif、I-box	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS	Dasgupta et al.,2020
	CitWAX	HSE、Skn-1 motif、I-box、TCA element	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	CitJuSac	Box4、HSE、ABRE、Box1	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	CitVO1	GARE-motif、Box1、HSE、ARE、ABER	柑橘 Citrus reticulata	番茄 Solanum lycopersicum	GUS
	PamMybAp	ARE、HSE、BOx4、as-2-box、MBS	李子 Prunus salicina	番茄 Solanum lycopersicum	GUS
	GSX7R	ACE、Box4、CAT box、G-Box、G-box、GATA motif、GC motif、I-box、Sp1、chs-unit1 M1	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Lin et al.,2022
	AraSSU	/	拟南芥 Arabidopsis thaliana	鹰嘴豆 Cicer arietinum	cry2Aa	Boruah et al.,2023
	BiGSSP2	POsrbcs-62、OsAct1、POsrbcs-550	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS	Bai et al.,2020
	BiGSSP3	POsrbcs-62、OsAct1、4×GEAT	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
	BiGSSP7	POsrbcs-550、PD540-544、OsAct1、OsTub6I	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
	BiGSSP6	OsAct1	人工合成 Synthetic	水稻 Oryza sativa	GFP、GUS
维管束特异性启动子 Vascular tissue-specific promoter	DROT1	/	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Sun et al.,2022
	SUC2	/	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Archer et al.,2023
	GmTIP1;6	MBS、ABRE、CGTCA-motif、TGACG-motif、TATC-box、TGA	毛豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Feng et al.,2022

激素Hormone	来源基因 Source gene	顺式作用元件cis-Acting element		来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
激素Hormone	来源基因 Source gene	名称 Name	作用 Function	来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
IAA	NtCel7	CATATG-motif	IAA反应元件IAA response element	烟草 Nicotiana tabacum	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2007a
		TATCCA-motif	GA反应元件GA response element
		W-box	SA、病原反应元件SA and pathogen response elements
		GT-1	病原体、盐胁迫pathogen、salt stress
	KnLBD1	RY-motif、CCTG-motif	IAA反应元件IAA response element	丝状绿藻 K. nitens	烟草 Nicotiana tabacum	GUS	Tounosu et al.,2023
	Dof5.8	ARF-bs	IAA反应因子结合位点IAA response factor binding site	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Konishi et al.,2015
	SlGH3.4	TGTCAC-motif	IAA反应元件IAA response element	番茄 Solanum lycopersicum	烟草 Nicotiana tabacum	GUS	Liao et al.,2015
	PIN1、PIN3、PIN7	AAGACA-motif、GAGACA-motif、DR5、GGGACA-motif	IAA反应元件 IAA response element	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Krogan et al.,2016
	LRP1	ARF7-bs、ARF9-bs	IAA反应因子结合位点IAA response factor binding site	拟南芥Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Singh et al., 2020
	10x2DR5-2DR5v2	DR5、DR5v2	IAA反应元件IAA response element	人工合成 Synthetic	拟南芥 Arabidopsis thaliana	GUS	Fernandez-Moreno et al.,2024
ABA	ADF5	ACGT/C	ABA响应元件核心基ABA-responsive element core motif	拟南芥Arabidopsis thaliana	烟草 Nicotiana tabacum	LUC	Qian et al.,2019
	Rab16A	G-box、Motif I	ABA响应元件 ABA response element	水稻 Oryza sativa	水稻 Oryza sativa	LUC	Song et al.,2020
	OsPSY3	G-box	ABA响应元件 ABA response element	水稻 Oryza sativa	/	/	Welsch et al.,2008
	OsPSY3	CE	干旱、盐胁迫响应元件 Drought and salt stress response elements	水稻 Oryza sativa	/	/	Welsch et al.,2008
	HvSnRK1	ABRE、LRE	ABA响应元件、光反应元件 ABA response elements、light response elements	大麦 Hordeum vulgare	/	/	Chen et al.,2021c
	SynP16	ACGTGTC-motif	like-ABA响应元件ABA-like response elements	人工合成 Synthetic	大豆 Glycine max	GUS	Jameel et al.,2020
		TACGTGGC-motif	ABA响应元件ABA response element
		GCC-box	ETH响应元件ETH response element
		Root--motif	根特异性元件root-specific elements
	GmRD26	CCACGTGG-motif、TCACGTGGC-motif、G-box、	ABA响应元件ABA response element	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Freitas et al.,2019
	GmRD26	DREB、MYC、MYB	干旱响应元件Drought response element	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Freitas et al.,2019
	AtNCED3	GGCACGTG-motif、GCCACGTC-motif、CACGTGGC-motif	ABA响应元件ABA response element	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Yang & Tan,2014
	PSCBV-YZ2060	TACGTG-motif、GCCACGT-motif	ABA响应元件ABA response element	甘蔗花叶病毒 SCMV	甘蔗 Saccharum officinarum	GUS	Sun et al.,2024
	osr40c1	GACAC-motif、G-box、GAGGC-motif、	ABA响应元件ABA response element	水稻 Oryza sativa	/	/	de Silva et al.,2017
		AAACAGA-motif	GA响应元件GA response element
		ATCT-motif、TGA-element、BoxI	光响应元件Light response elements
		Skn-1 motif	胚乳特异性表达Endosperm-specific expression
		Box E、Box-W1	真菌诱导Fungal induction
GA	Amy32b	GTAACAGAGTCTGG-motif	GA响应元件GA response elements	大麦02SHordeum vulgare	大麦02SHordeum vulgare	GUS	Rogers & Rogers,1992
	Amy32b	02S	耦合元件Coupling elements	大麦02SHordeum vulgare	大麦02SHordeum vulgare	GUS	Rogers & Rogers,1992
	GhGlcAT1	MYB1AT、MYC	MYB 结合基序MYB binding motif、干旱响应元件Drought response elements	陆地棉Gossypium hirsutum	陆地棉Gossypium hirsutum	GUS	Yadav et al.,2017
	Rep1	TAACAGA-motif、CAACTCCC-motif、CAACTACAC-motif、AAGACAAT-motif	GA响应元件GA response elements	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Sutoh &Yamauchi,2003
	GmGASA32	/	/	大豆 Glycine max	大豆 Glycine max	GUS	Chen et al.,2021a
ETH	E4、E8	/	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Kneissl & Deikman,1996；Xu et al.,1996
	10x2EBS-S10	2EBS-S10	/	合成 Synthetic	拟南芥 Arabidopsis thaliana	GUS	Fernandez-Moreno et al.,2024
	CmaACS	ERE、W-box	ETH反应元件ETH response elements	南瓜 Cucurbita pepo	/	/	Wang et al.,2022
		TCA-element	SA反应元件SA response elements
ETH	CmaACS	ABRE	ABA反应元件ABA response elements	南瓜 Cucurbita pepo	/	/	Wang et al.,2022
		GARE	GA反应元件GA response elements
		ARE、LTR、MBS	应激反应元件Stress response elements
	OsTAR2	EBS-element	ETH响应元件ETH response elements	水稻 Oryza sativa	烟草、水稻 Nicotiana tabacum，Oryza sativa	LUC、GUS	Zhou et al.,2022
SA	SFR2	W-box	SA诱导元件SA-inducible elements	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Rocher et al.,2005
SA	FUASCsV8CP	/	/	合成 Synthetic	烟草 Nicotiana tabacum	GUS	Deb & Dey,2019
	Cyr1P4、Cyr1P5	TCA-element、WRKY71OS、WBOXATNPR1	SA诱导元件SA-inducible elements	黄花叶病毒 Cucumber mosaic virus	烟草、菠菜、洋葱、绿豆 Nicotiana tabacum，Spinacia oleracea，Allium cepa，Vigna radiata	GUS	Maiti et al.,2015
		MYB1AT、MYBCORE、MYCCONSENSUSA	响应干旱、ABA诱导Responsive to drought and ABA induction
		TAAG-motif、YACTT	叶肉细胞特异性元件Mesophyll-specific elements
	CmYLCV	as-1、W-box1	SA诱导顺式元件SA-inducible cis-elements	夜来香黄叶卷曲病毒 MJCMV	烟草、拟南芥 Nicotiana tabacum，Arabidopsis thaliana	GUS	Sarkar et al.,2018
	VqMYB15	MRE、Box Ⅰ、Box4、LAMP-element	光敏元件Photosensitive elements	葡萄 Vitis vinifera	烟草、葡萄 Nicotiana tabacum，Vitis vinifera	GUS	Li et al.,2020b
		TATTTTTCTC-motif、CCATCTATTT-motif、GAGAAGAAGA-motif	SA反应元件 SA response elements
		ABRE、MYC、CAATTG-motif	干旱、ABA诱导元件Drought and ABA-inducible elements
	D100	DRE core、GT1-box、MYC、MYCR、as-1	响应环境胁迫Responsive to environmental stress	大丽花花叶病毒 DMV	烟草 Nicotiana tabacum	GUS	Khadanga et al.,2021
	FvTrihelix6	G-box、ARE、MBS、LTR	应激响应元件 Stress response elements	草莓 Fragaria × ananassa	烟草 Nicotiana tabacum	GUS	Fan et al.,2023
	FvTrihelix6	ABRE	ABA响应元件 ABA response element	草莓 Fragaria × ananassa	烟草 Nicotiana tabacum	GUS	Fan et al.,2023
	MiWRKY53	W-box、ERE	ETH反应元件 ETH response element	桑树 Morus alba	拟南芥 Arabidopsis thaliana	GUS	Negi & Khurana,2021
		TCA-element	SA响应元件 SA response element
		ABRERATCAL、ACGTG-motif、TACGTG-motif、ACCGAC-motif	ABA响应元件 ABA response element
		TGA-element	IAA响应元件 IAA response element
		MYC	干旱响应元件 Drought response elements
JA	CYP81D11	TGACGTCA-motif	结合TGA转录因子Binding to TGA transcription factors	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Köster et al.,2012
	CYP81D11	as-1、G-box	结合JA调节的转录因子Transcription factors regulated by JA binding	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Köster et al.,2012
	TSA1	/	/	拟南芥 Arabidopsis thaliana	拟南芥、烟草 Arabidopsis thaliana，Nicotiana tabacum	GFP	Geem et al.,2019
	Zmap	CGTCA-motif、TGACG-motif	MeJA响应元件 MeJA response elements	玉米 Zea mays	烟草 Nicotiana tabacum	GUS	Jin et al.,2019
		GGTCCAT-motif	IAA响应元件 IAA response elements
		Box-II、GA-motif	光响应元件 Light response elements
		H-box、MBS	ABA、干旱响应元件ABA and drought response elements
	VvPR1	ACGTG-motif、MBS	ABA、干旱响应响应元件 ABA and drought response elements	葡萄 Vitis vinifera	烟草 Nicotiana tabacum	GUS	Rahman et al.,2022b
		ERE	ETH响应元件					ETH and drought response elements
		P-box	GA响应元件 GA response elements
		TCA	SA反应元件 SA response elements
		CGTCA-motif	MeJA响应元件 MeJA response elements
		LS7、G-box、AT1-motif	光响应元件 Light response elements
	TkSRPP	CGTCA-motif	MeJA响应元件 MeJA response elements	蒲公英 Taraxacum officinale	烟草 Nicotiana tabacum	GUS	Dong et al.,2023
		TGA-element	IAA响应元件 IAA response elements
		W-box、TTGAC-motif	SA响应元件 SA response elements
		CANNTG-motif、CNGTTR-motif、ACTG-motif	ABA响应元件 GA response elements
		MBS	干旱响应 Drought response

激素Hormone	来源基因 Source gene	顺式作用元件cis-Acting element		来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
激素Hormone	来源基因 Source gene	名称 Name	作用 Function	来源物种 Source species	转化物种 Transformed species	报告基因 Reporter gene	参考文献 Reference
IAA	NtCel7	CATATG-motif	IAA反应元件IAA response element	烟草 Nicotiana tabacum	拟南芥 Arabidopsis thaliana	GUS	Wang et al.,2007a
		TATCCA-motif	GA反应元件GA response element
		W-box	SA、病原反应元件SA and pathogen response elements
		GT-1	病原体、盐胁迫pathogen、salt stress
	KnLBD1	RY-motif、CCTG-motif	IAA反应元件IAA response element	丝状绿藻 K. nitens	烟草 Nicotiana tabacum	GUS	Tounosu et al.,2023
	Dof5.8	ARF-bs	IAA反应因子结合位点IAA response factor binding site	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Konishi et al.,2015
	SlGH3.4	TGTCAC-motif	IAA反应元件IAA response element	番茄 Solanum lycopersicum	烟草 Nicotiana tabacum	GUS	Liao et al.,2015
	PIN1、PIN3、PIN7	AAGACA-motif、GAGACA-motif、DR5、GGGACA-motif	IAA反应元件 IAA response element	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Krogan et al.,2016
	LRP1	ARF7-bs、ARF9-bs	IAA反应因子结合位点IAA response factor binding site	拟南芥Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Singh et al., 2020
	10x2DR5-2DR5v2	DR5、DR5v2	IAA反应元件IAA response element	人工合成 Synthetic	拟南芥 Arabidopsis thaliana	GUS	Fernandez-Moreno et al.,2024
ABA	ADF5	ACGT/C	ABA响应元件核心基ABA-responsive element core motif	拟南芥Arabidopsis thaliana	烟草 Nicotiana tabacum	LUC	Qian et al.,2019
	Rab16A	G-box、Motif I	ABA响应元件 ABA response element	水稻 Oryza sativa	水稻 Oryza sativa	LUC	Song et al.,2020
	OsPSY3	G-box	ABA响应元件 ABA response element	水稻 Oryza sativa	/	/	Welsch et al.,2008
	OsPSY3	CE	干旱、盐胁迫响应元件 Drought and salt stress response elements	水稻 Oryza sativa	/	/	Welsch et al.,2008
	HvSnRK1	ABRE、LRE	ABA响应元件、光反应元件 ABA response elements、light response elements	大麦 Hordeum vulgare	/	/	Chen et al.,2021c
	SynP16	ACGTGTC-motif	like-ABA响应元件ABA-like response elements	人工合成 Synthetic	大豆 Glycine max	GUS	Jameel et al.,2020
		TACGTGGC-motif	ABA响应元件ABA response element
		GCC-box	ETH响应元件ETH response element
		Root--motif	根特异性元件root-specific elements
	GmRD26	CCACGTGG-motif、TCACGTGGC-motif、G-box、	ABA响应元件ABA response element	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Freitas et al.,2019
	GmRD26	DREB、MYC、MYB	干旱响应元件Drought response element	大豆 Glycine max	拟南芥 Arabidopsis thaliana	GUS	Freitas et al.,2019
	AtNCED3	GGCACGTG-motif、GCCACGTC-motif、CACGTGGC-motif	ABA响应元件ABA response element	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Yang & Tan,2014
	PSCBV-YZ2060	TACGTG-motif、GCCACGT-motif	ABA响应元件ABA response element	甘蔗花叶病毒 SCMV	甘蔗 Saccharum officinarum	GUS	Sun et al.,2024
	osr40c1	GACAC-motif、G-box、GAGGC-motif、	ABA响应元件ABA response element	水稻 Oryza sativa	/	/	de Silva et al.,2017
		AAACAGA-motif	GA响应元件GA response element
		ATCT-motif、TGA-element、BoxI	光响应元件Light response elements
		Skn-1 motif	胚乳特异性表达Endosperm-specific expression
		Box E、Box-W1	真菌诱导Fungal induction
GA	Amy32b	GTAACAGAGTCTGG-motif	GA响应元件GA response elements	大麦02SHordeum vulgare	大麦02SHordeum vulgare	GUS	Rogers & Rogers,1992
	Amy32b	02S	耦合元件Coupling elements	大麦02SHordeum vulgare	大麦02SHordeum vulgare	GUS	Rogers & Rogers,1992
	GhGlcAT1	MYB1AT、MYC	MYB 结合基序MYB binding motif、干旱响应元件Drought response elements	陆地棉Gossypium hirsutum	陆地棉Gossypium hirsutum	GUS	Yadav et al.,2017
	Rep1	TAACAGA-motif、CAACTCCC-motif、CAACTACAC-motif、AAGACAAT-motif	GA响应元件GA response elements	水稻 Oryza sativa	水稻 Oryza sativa	GUS	Sutoh &Yamauchi,2003
	GmGASA32	/	/	大豆 Glycine max	大豆 Glycine max	GUS	Chen et al.,2021a
ETH	E4、E8	/	/	番茄 Solanum lycopersicum	番茄 Solanum lycopersicum	GUS	Kneissl & Deikman,1996；Xu et al.,1996
	10x2EBS-S10	2EBS-S10	/	合成 Synthetic	拟南芥 Arabidopsis thaliana	GUS	Fernandez-Moreno et al.,2024
	CmaACS	ERE、W-box	ETH反应元件ETH response elements	南瓜 Cucurbita pepo	/	/	Wang et al.,2022
		TCA-element	SA反应元件SA response elements
ETH	CmaACS	ABRE	ABA反应元件ABA response elements	南瓜 Cucurbita pepo	/	/	Wang et al.,2022
		GARE	GA反应元件GA response elements
		ARE、LTR、MBS	应激反应元件Stress response elements
	OsTAR2	EBS-element	ETH响应元件ETH response elements	水稻 Oryza sativa	烟草、水稻 Nicotiana tabacum，Oryza sativa	LUC、GUS	Zhou et al.,2022
SA	SFR2	W-box	SA诱导元件SA-inducible elements	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Rocher et al.,2005
SA	FUASCsV8CP	/	/	合成 Synthetic	烟草 Nicotiana tabacum	GUS	Deb & Dey,2019
	Cyr1P4、Cyr1P5	TCA-element、WRKY71OS、WBOXATNPR1	SA诱导元件SA-inducible elements	黄花叶病毒 Cucumber mosaic virus	烟草、菠菜、洋葱、绿豆 Nicotiana tabacum，Spinacia oleracea，Allium cepa，Vigna radiata	GUS	Maiti et al.,2015
		MYB1AT、MYBCORE、MYCCONSENSUSA	响应干旱、ABA诱导Responsive to drought and ABA induction
		TAAG-motif、YACTT	叶肉细胞特异性元件Mesophyll-specific elements
	CmYLCV	as-1、W-box1	SA诱导顺式元件SA-inducible cis-elements	夜来香黄叶卷曲病毒 MJCMV	烟草、拟南芥 Nicotiana tabacum，Arabidopsis thaliana	GUS	Sarkar et al.,2018
	VqMYB15	MRE、Box Ⅰ、Box4、LAMP-element	光敏元件Photosensitive elements	葡萄 Vitis vinifera	烟草、葡萄 Nicotiana tabacum，Vitis vinifera	GUS	Li et al.,2020b
		TATTTTTCTC-motif、CCATCTATTT-motif、GAGAAGAAGA-motif	SA反应元件 SA response elements
		ABRE、MYC、CAATTG-motif	干旱、ABA诱导元件Drought and ABA-inducible elements
	D100	DRE core、GT1-box、MYC、MYCR、as-1	响应环境胁迫Responsive to environmental stress	大丽花花叶病毒 DMV	烟草 Nicotiana tabacum	GUS	Khadanga et al.,2021
	FvTrihelix6	G-box、ARE、MBS、LTR	应激响应元件 Stress response elements	草莓 Fragaria × ananassa	烟草 Nicotiana tabacum	GUS	Fan et al.,2023
	FvTrihelix6	ABRE	ABA响应元件 ABA response element	草莓 Fragaria × ananassa	烟草 Nicotiana tabacum	GUS	Fan et al.,2023
	MiWRKY53	W-box、ERE	ETH反应元件 ETH response element	桑树 Morus alba	拟南芥 Arabidopsis thaliana	GUS	Negi & Khurana,2021
		TCA-element	SA响应元件 SA response element
		ABRERATCAL、ACGTG-motif、TACGTG-motif、ACCGAC-motif	ABA响应元件 ABA response element
		TGA-element	IAA响应元件 IAA response element
		MYC	干旱响应元件 Drought response elements
JA	CYP81D11	TGACGTCA-motif	结合TGA转录因子Binding to TGA transcription factors	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Köster et al.,2012
	CYP81D11	as-1、G-box	结合JA调节的转录因子Transcription factors regulated by JA binding	拟南芥 Arabidopsis thaliana	拟南芥 Arabidopsis thaliana	GUS	Köster et al.,2012
	TSA1	/	/	拟南芥 Arabidopsis thaliana	拟南芥、烟草 Arabidopsis thaliana，Nicotiana tabacum	GFP	Geem et al.,2019
	Zmap	CGTCA-motif、TGACG-motif	MeJA响应元件 MeJA response elements	玉米 Zea mays	烟草 Nicotiana tabacum	GUS	Jin et al.,2019
		GGTCCAT-motif	IAA响应元件 IAA response elements
		Box-II、GA-motif	光响应元件 Light response elements
		H-box、MBS	ABA、干旱响应元件ABA and drought response elements
	VvPR1	ACGTG-motif、MBS	ABA、干旱响应响应元件 ABA and drought response elements	葡萄 Vitis vinifera	烟草 Nicotiana tabacum	GUS	Rahman et al.,2022b
		ERE	ETH响应元件					ETH and drought response elements
		P-box	GA响应元件 GA response elements
		TCA	SA反应元件 SA response elements
		CGTCA-motif	MeJA响应元件 MeJA response elements
		LS7、G-box、AT1-motif	光响应元件 Light response elements
	TkSRPP	CGTCA-motif	MeJA响应元件 MeJA response elements	蒲公英 Taraxacum officinale	烟草 Nicotiana tabacum	GUS	Dong et al.,2023
		TGA-element	IAA响应元件 IAA response elements
		W-box、TTGAC-motif	SA响应元件 SA response elements
		CANNTG-motif、CNGTTR-motif、ACTG-motif	ABA响应元件 GA response elements
		MBS	干旱响应 Drought response