花青素转录因子调控机制及代谢工程研究进展

doi:10.11983/CBB18016

摘要/Abstract

摘要： 花青素是广泛存在于植物中的一类重要的类黄酮化合物, 在植物生长发育和人类营养保健方面具有重要价值。花青素的生物合成途径已经解析得比较清楚, 但花青素的代谢调控网络还在不断完善。调控花青素生物合成的转录因子主要包括MYB、bHLH和WD40三大类, 这些转录因子通过激活或抑制CHS、ANS和DFR等花青素途径关键结构基因的表达水平, 进而决定花青素积累的部位与水平。该文结合国内外花青素生物合成与转录调控方面的研究进展, 简要介绍了花青素的生物合成途径, 归纳总结了模式植物中花青素代谢调控的分子机理, 尤其是MYB、bHLH和WD40三类主要转录因子的调控机理, 以及这些转录因子在观赏植物和水果等经济作物花青素代谢工程中的应用。该文将为系统阐明花青素的转录调控机制和利用代谢工程改良花青素的相关研究提供有益参考。

关键词: 花青素, 转录因子, 转录调控, 代谢工程

Abstract: Anthocyanins are among the most important flavonoid compounds widely present in plants. Anthocyanins play significant roles in plant growth and development as well as human nutrition and health care. The anthocyanin biosynthetic pathway has been widely documented, and the anthocyanin metabolic regulation network is being constantly improved. The transcription factors that regulate anthocyanin biosynthesis mainly include three classes: MYB, bHLH and WD40 proteins. The proteins regulate the accumulation, location and levels of anthocyanins by activating or suppressing the expression of key structural genes, including CHS, ANS and DFR. This review briefly introduces the anthocyanin biosynthetic pathway and summarizes the molecular mechanism of transcriptional regulation based on recent progress. It mainly focuses on the molecular mechanism of MYB, bHLH and WD40 transcription factors in the regulation of anthocyanins in model plants. In addition, it summarizes the use of these transcription factors in anthocyanin metabolic engineering in ornamental plants and fruit crops. This review will provide valuable references for the in-depth investigation of transcriptional regulation and improving anthocyanins by metabolic engineering.

Key words: anthocyanins, transcription factor, transcriptional regulation, metabolic engineering

宋雪薇,魏解冰,狄少康,庞永珍. 花青素转录因子调控机制及代谢工程研究进展. 植物学报, 2019, 54(1): 133-156.

Xuewei Song,Jiebing Wei,Shaokang Di,Yongzhen Pang. Recent Advances in the Regulation Mechanism of Transcription Factors and Metabolic Engineering of Anthocyanins. Chinese Bulletin of Botany, 2019, 54(1): 133-156.

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https://www.chinbullbotany.com/CN/Y2019/V54/I1/133

图/表 5

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物种名称	拉丁名	基因名称	类型	参考文献
玉米	Zea mays	C1	激活	Cone et al., 1986
		PL	激活	Cone et al., 1993a
		PL-BH	激活	Cone et al., 1993b
拟南芥	Arabidopsis thaliana	AtCPC	抑制	Zhang et al., 2009
		AtMYB60	抑制	Park et al., 2008
		AtMYBL2	抑制	Matsui et al., 2008
		AtMYB75/PAP1	激活	Borevitz et al., 2000
		AtMYB90/PAP2	激活	Borevitz et al., 2000
		AtMYB113	激活	Gonzalez et al., 2008
		AtMYB114	激活	Gonzalez et al., 2008
		AtMYB56	激活	Jeong et al., 2018
蒺藜苜蓿	Medicago truncatula	LAP1	激活	Peel et al., 2009
矮牵牛	Petunia hybrida	AN2	激活	Quattrocchio et al., 1999
		DPL	激活	Albert et al., 2011
		PHZ	激活	Albert et al., 2011
		MYBx	抑制	Albert et al., 2014
		MYB27	抑制	Albert et al., 2014
金鱼草	Antirrhinum majus	ROSEA1	激活	Schwinn et al., 2006
		ROSEA2	激活	Schwinn et al., 2006
		VENOSA	激活	Schwinn et al., 2016
烟草	Nicotiana tabacum	NtAN2	激活	Pattanaik et al., 2010
苹果	Malus domestica	MdMYB1	激活	Takos et al., 2006
		MdMYB3	激活	Vimolmangkang et al., 2013
		MdMYB6	激活	Gao et al., 2011
		MdMYB10	激活	Espley et al., 2007
		MdMYB110a	激活	Chagné et al., 2013
		MdMYB9	激活	An et al., 2014
		MdMYB11	激活	An et al., 2014
		MdMYBA	激活	Ban et al., 2007
		MdMYB16	抑制	Wang et al., 2010
葡萄	Vitis labruscana	VlMYBA1-1	激活	Kobayashi et al., 2002
		VlMYBA1-2	激活	Cutanda-Perez et al., 2009
		VlMYBA1-3	激活	Azuma et al., 2008
		VlMYBA2	激活	Geekiyanage et al., 2007
	V. vinifera	VvMYB5a	激活	Deluc et al., 2006
		VvMYB5b	激活	Deluc et al., 2008
		VvMYBA1	激活	Kobayashi et al., 2004
		VvMYBA2	激活	Kobayashi et al., 2004
		VvMYBPA1	激活	Passeri et al., 2017
		VvMYBC2-L1	抑制	Cavallini et al., 2015
		VvMYBC2-L3	抑制	Cavallini et al., 2015
西洋梨	Pyrus communis	PcMYB10	激活	Pierantoni et al., 2010
沙梨	P. pyrifolia	PyMYB10	激活	Feng et al., 2010
		PyMYB114	激活	Yao et al., 2017a
物种名称	拉丁名	基因名称	类型	参考文献
	P. communis	PcMYB10	激活	Wang et al., 2013
山竹	Garcinia mangostana	GmMYB10	激活	Palapol et al., 2009
甜樱桃	Prunus avium	PacMYBA	激活	Shen et al., 2014
		PaMYB10	激活	Starkevi? et al., 2015
荔枝	Litchi chinensis	LcMYB1	激活	Lai et al., 2014
油桃	Prunus persica	PpMYB10	激活	Ravaglia et al., 2013
甜橙	Citrus sinensis	CsRUBY	激活	Butelli et al., 2012
杨梅	Myrica rubra	MrMYB1	激活	Niu et al., 2010
草莓	Fragaria ananassa	FaMYB10	激活	Medina-Puche et al., 2014
		FaMYB1	抑制	Aharoni et al., 2001
猕猴桃	Actinidia chinensis	MYB110a	激活	Fraser et al., 2013
亚洲杂交百合	Lilium spp.	LhMYB6	激活	Yamagishi et al., 2010
		LhMYB12	激活	Yamagishi et al., 2010
		LhMYB12-Lat	激活	Yamagishi et al., 2014
非洲菊	Gerbera hybrida	GMYB10	激活	Elomaa et al., 2003
菊花	Chrysanthemum morifolium	CmMYB6	激活	Liu et al., 2015
文心兰	Oncidium gower	OgMYB1	激活	Chiou et al., 2008
龙胆	Gentian triflora	GtMYB3	激活	Nakatsuka et al., 2008
紫苏	Perilla frutescens	Myb-p1	激活	Gong et al., 1999
番茄	Lycopersicon esculentum	LeANT1	激活	Mathews et al., 2003
红薯	Ipomoea batatas	IbMYB1	激活	Mano et al., 2007
马铃薯	Solanum tuberosum	StAN1	激活	Jung et al., 2009
		StAN2	激活	Jung et al., 2009
		StMTF1	激活	Rommens et al., 2008
结球甘蓝	Brassica oleracea var. capitata	BoMYB2	激活	Yuan et al., 2009
花椰菜	B. oleracea var. botrytis	PURPLE	激活	Chiu and Yeh, 2010
紫背天葵	Gynura bicolor	GbMYB1	激活	Shimizu et al., 2011
羽衣甘蓝	B. oeracea var. acephala	BoPAP1	激活	Zhang et al., 2012
淫羊藿	Epimedium sagittatum	EsMYBA1	激活	Huang et al., 2013a
		EsMYB9	激活	Huang et al., 2017
胡萝卜	Raphanus sativus	RsMYB1	激活	Lim et al., 2016
洋葱	Allium cepa	MYB1	激活	Schwinn et al., 2016
小麦	Triticum aestivum	TaPL1	激活	Shin et al., 2016
		TaMYB3	激活	Li et al., 2017
红掌	Anthurium andraeanum	AaMYB2	激活	Li et al., 2016a
大豆	Glycine max	GmMYB-G20-1	激活	Takahashi et al., 2011
梅	Prunus mume	PmMYBa1	激活	Zhang et al., 2017
葡萄风信子	Muscari armeniacum	MaAN2	激活	Chen et al., 2017
智利草莓	Fragaria chiloensis	FcMYB1	抑制	Salvatierra et al., 2013
芜菁	B. rapa	BrMYB4	抑制	Zhang et al., 2014a
银杏	Ginkgo biloba	GbMYBF2	抑制	Xu et al., 2014a
		GbMYBFL	激活	Zhang et al., 2018
杨树	Populus trichocarpa	PtrRML1	抑制	Hu et al., 2016

物种名称	拉丁名	基因名称	类型	参考文献
玉米	Zea mays	C1	激活	Cone et al., 1986
		PL	激活	Cone et al., 1993a
		PL-BH	激活	Cone et al., 1993b
拟南芥	Arabidopsis thaliana	AtCPC	抑制	Zhang et al., 2009
		AtMYB60	抑制	Park et al., 2008
		AtMYBL2	抑制	Matsui et al., 2008
		AtMYB75/PAP1	激活	Borevitz et al., 2000
		AtMYB90/PAP2	激活	Borevitz et al., 2000
		AtMYB113	激活	Gonzalez et al., 2008
		AtMYB114	激活	Gonzalez et al., 2008
		AtMYB56	激活	Jeong et al., 2018
蒺藜苜蓿	Medicago truncatula	LAP1	激活	Peel et al., 2009
矮牵牛	Petunia hybrida	AN2	激活	Quattrocchio et al., 1999
		DPL	激活	Albert et al., 2011
		PHZ	激活	Albert et al., 2011
		MYBx	抑制	Albert et al., 2014
		MYB27	抑制	Albert et al., 2014
金鱼草	Antirrhinum majus	ROSEA1	激活	Schwinn et al., 2006
		ROSEA2	激活	Schwinn et al., 2006
		VENOSA	激活	Schwinn et al., 2016
烟草	Nicotiana tabacum	NtAN2	激活	Pattanaik et al., 2010
苹果	Malus domestica	MdMYB1	激活	Takos et al., 2006
		MdMYB3	激活	Vimolmangkang et al., 2013
		MdMYB6	激活	Gao et al., 2011
		MdMYB10	激活	Espley et al., 2007
		MdMYB110a	激活	Chagné et al., 2013
		MdMYB9	激活	An et al., 2014
		MdMYB11	激活	An et al., 2014
		MdMYBA	激活	Ban et al., 2007
		MdMYB16	抑制	Wang et al., 2010
葡萄	Vitis labruscana	VlMYBA1-1	激活	Kobayashi et al., 2002
		VlMYBA1-2	激活	Cutanda-Perez et al., 2009
		VlMYBA1-3	激活	Azuma et al., 2008
		VlMYBA2	激活	Geekiyanage et al., 2007
	V. vinifera	VvMYB5a	激活	Deluc et al., 2006
		VvMYB5b	激活	Deluc et al., 2008
		VvMYBA1	激活	Kobayashi et al., 2004
		VvMYBA2	激活	Kobayashi et al., 2004
		VvMYBPA1	激活	Passeri et al., 2017
		VvMYBC2-L1	抑制	Cavallini et al., 2015
		VvMYBC2-L3	抑制	Cavallini et al., 2015
西洋梨	Pyrus communis	PcMYB10	激活	Pierantoni et al., 2010
沙梨	P. pyrifolia	PyMYB10	激活	Feng et al., 2010
		PyMYB114	激活	Yao et al., 2017a
物种名称	拉丁名	基因名称	类型	参考文献
	P. communis	PcMYB10	激活	Wang et al., 2013
山竹	Garcinia mangostana	GmMYB10	激活	Palapol et al., 2009
甜樱桃	Prunus avium	PacMYBA	激活	Shen et al., 2014
		PaMYB10	激活	Starkevi? et al., 2015
荔枝	Litchi chinensis	LcMYB1	激活	Lai et al., 2014
油桃	Prunus persica	PpMYB10	激活	Ravaglia et al., 2013
甜橙	Citrus sinensis	CsRUBY	激活	Butelli et al., 2012
杨梅	Myrica rubra	MrMYB1	激活	Niu et al., 2010
草莓	Fragaria ananassa	FaMYB10	激活	Medina-Puche et al., 2014
		FaMYB1	抑制	Aharoni et al., 2001
猕猴桃	Actinidia chinensis	MYB110a	激活	Fraser et al., 2013
亚洲杂交百合	Lilium spp.	LhMYB6	激活	Yamagishi et al., 2010
		LhMYB12	激活	Yamagishi et al., 2010
		LhMYB12-Lat	激活	Yamagishi et al., 2014
非洲菊	Gerbera hybrida	GMYB10	激活	Elomaa et al., 2003
菊花	Chrysanthemum morifolium	CmMYB6	激活	Liu et al., 2015
文心兰	Oncidium gower	OgMYB1	激活	Chiou et al., 2008
龙胆	Gentian triflora	GtMYB3	激活	Nakatsuka et al., 2008
紫苏	Perilla frutescens	Myb-p1	激活	Gong et al., 1999
番茄	Lycopersicon esculentum	LeANT1	激活	Mathews et al., 2003
红薯	Ipomoea batatas	IbMYB1	激活	Mano et al., 2007
马铃薯	Solanum tuberosum	StAN1	激活	Jung et al., 2009
		StAN2	激活	Jung et al., 2009
		StMTF1	激活	Rommens et al., 2008
结球甘蓝	Brassica oleracea var. capitata	BoMYB2	激活	Yuan et al., 2009
花椰菜	B. oleracea var. botrytis	PURPLE	激活	Chiu and Yeh, 2010
紫背天葵	Gynura bicolor	GbMYB1	激活	Shimizu et al., 2011
羽衣甘蓝	B. oeracea var. acephala	BoPAP1	激活	Zhang et al., 2012
淫羊藿	Epimedium sagittatum	EsMYBA1	激活	Huang et al., 2013a
		EsMYB9	激活	Huang et al., 2017
胡萝卜	Raphanus sativus	RsMYB1	激活	Lim et al., 2016
洋葱	Allium cepa	MYB1	激活	Schwinn et al., 2016
小麦	Triticum aestivum	TaPL1	激活	Shin et al., 2016
		TaMYB3	激活	Li et al., 2017
红掌	Anthurium andraeanum	AaMYB2	激活	Li et al., 2016a
大豆	Glycine max	GmMYB-G20-1	激活	Takahashi et al., 2011
梅	Prunus mume	PmMYBa1	激活	Zhang et al., 2017
葡萄风信子	Muscari armeniacum	MaAN2	激活	Chen et al., 2017
智利草莓	Fragaria chiloensis	FcMYB1	抑制	Salvatierra et al., 2013
芜菁	B. rapa	BrMYB4	抑制	Zhang et al., 2014a
银杏	Ginkgo biloba	GbMYBF2	抑制	Xu et al., 2014a
		GbMYBFL	激活	Zhang et al., 2018
杨树	Populus trichocarpa	PtrRML1	抑制	Hu et al., 2016

物种名称	拉丁名	基因名称	类型	参考文献
玉米	Zea mays	R1	激活	Chandler et al., 1989
		Lc	激活	Ludwig et al., 1989
		B1	激活	Chandler et al., 1989
		Sn	激活	Tonelli et al., 1991
金鱼草	Antirrhinum majus	DELILA	激活	Martin et al.,1991
矮牵牛	Petunia hybrida	JAF13	激活	Spelt et al., 2000
		AN1	激活	Spelt et al., 2000
拟南芥	Arabidopsis thaliana	GL3	激活	Payne et al., 2000
		EGL3	激活	Zhang et al., 2003
		MYC146	激活	Ramsay et al., 2003
		TT8	激活	Nesi et al., 2000
蒺藜苜蓿	Medicago truncatula	MtTT8	激活	Li et al., 2016b
苹果	Malus domestica	MdbHLH3	激活	Xie et al., 2012
		MdbHLH33	激活	Espley et al., 2007
		MdMYC2	激活	An et al., 2016
烟草	Nicotiana tabacum	NtAn1a	激活	Bai et al., 2011
		NtAn1b	激活	Bai et al., 2011
菊花	Chrysanthemum morifolium	CmbHLH	激活	Xiang et al., 2015
非洲菊	Gerbera hybrida	GMYC1	激活	Elomaa et al., 2003
荔枝	Litchi chinensis	LcbHLH1	激活	Lai et al., 2016
		LcbHLH3	激活	Lai et al., 2016
龙胆	Gentian triflora	GtbHLH1	激活	Nakatsuka et al., 2008
杨梅	Myrica rubra	MrbHLH1	激活	Liu et al., 2013
花椰菜	Brassica oleracea var. botrytis	BobHLH1	激活	Chiu and Li, 2012
结球甘蓝	B. oleracea var. capitata	BoTT8	激活	Yuan et al., 2009
紫背天葵	Gynura bicolor	GbMYC1	激活	Shimizu et al., 2011
杂交百合	Lilium spp.	LhbHLH1	激活	Nakatsuka et al., 2009
番茄	Lycopersicon esculentum	AH	激活	Qiu et al., 2016
大丽花	Dahlia variabilis	DvIVS	抑制	Ohno et al., 2011
圆叶牵牛	Ipomoea purpurea	bHLH2	激活	Park et al., 2007
甜橙	Citrus sinensis	CsMYC2	激活	Cultrone et al., 2010
紫苏	Gynura bicolor	Myc-rp	激活	Gong et al., 1999
葡萄	Vitis vinifera	VvbHLH1	激活	Xu et al., 2014b
		VvMYC1	激活	Hichri et al., 2010
		VvMYCA1	激活	Hichri et al., 2010
		Myc-F3G1	激活	Yamazaki et al., 2003
甜樱桃	Prunus avium	PabHLH3	激活	Starkevi? et al., 2015
梨	Pyrus pyrifolia	PybHLH3	激活	Yao et al., 2017a
		LhbHLH2	激活	Nakatsuka et al., 2009
马铃薯	Solanum tuberosum	StbHLH1	激活	Payyavula et al., 2013
		StbHLH2	激活	Payyavula et al., 2013
		AH	激活	Qiu et al., 2016

物种名称	拉丁名	基因名称	类型	参考文献
玉米	Zea mays	R1	激活	Chandler et al., 1989
		Lc	激活	Ludwig et al., 1989
		B1	激活	Chandler et al., 1989
		Sn	激活	Tonelli et al., 1991
金鱼草	Antirrhinum majus	DELILA	激活	Martin et al.,1991
矮牵牛	Petunia hybrida	JAF13	激活	Spelt et al., 2000
		AN1	激活	Spelt et al., 2000
拟南芥	Arabidopsis thaliana	GL3	激活	Payne et al., 2000
		EGL3	激活	Zhang et al., 2003
		MYC146	激活	Ramsay et al., 2003
		TT8	激活	Nesi et al., 2000
蒺藜苜蓿	Medicago truncatula	MtTT8	激活	Li et al., 2016b
苹果	Malus domestica	MdbHLH3	激活	Xie et al., 2012
		MdbHLH33	激活	Espley et al., 2007
		MdMYC2	激活	An et al., 2016
烟草	Nicotiana tabacum	NtAn1a	激活	Bai et al., 2011
		NtAn1b	激活	Bai et al., 2011
菊花	Chrysanthemum morifolium	CmbHLH	激活	Xiang et al., 2015
非洲菊	Gerbera hybrida	GMYC1	激活	Elomaa et al., 2003
荔枝	Litchi chinensis	LcbHLH1	激活	Lai et al., 2016
		LcbHLH3	激活	Lai et al., 2016
龙胆	Gentian triflora	GtbHLH1	激活	Nakatsuka et al., 2008
杨梅	Myrica rubra	MrbHLH1	激活	Liu et al., 2013
花椰菜	Brassica oleracea var. botrytis	BobHLH1	激活	Chiu and Li, 2012
结球甘蓝	B. oleracea var. capitata	BoTT8	激活	Yuan et al., 2009
紫背天葵	Gynura bicolor	GbMYC1	激活	Shimizu et al., 2011
杂交百合	Lilium spp.	LhbHLH1	激活	Nakatsuka et al., 2009
番茄	Lycopersicon esculentum	AH	激活	Qiu et al., 2016
大丽花	Dahlia variabilis	DvIVS	抑制	Ohno et al., 2011
圆叶牵牛	Ipomoea purpurea	bHLH2	激活	Park et al., 2007
甜橙	Citrus sinensis	CsMYC2	激活	Cultrone et al., 2010
紫苏	Gynura bicolor	Myc-rp	激活	Gong et al., 1999
葡萄	Vitis vinifera	VvbHLH1	激活	Xu et al., 2014b
		VvMYC1	激活	Hichri et al., 2010
		VvMYCA1	激活	Hichri et al., 2010
		Myc-F3G1	激活	Yamazaki et al., 2003
甜樱桃	Prunus avium	PabHLH3	激活	Starkevi? et al., 2015
梨	Pyrus pyrifolia	PybHLH3	激活	Yao et al., 2017a
		LhbHLH2	激活	Nakatsuka et al., 2009
马铃薯	Solanum tuberosum	StbHLH1	激活	Payyavula et al., 2013
		StbHLH2	激活	Payyavula et al., 2013
		AH	激活	Qiu et al., 2016

物种名称	拉丁名	基因名称	类型	参考文献
矮牵牛	Petunia hybrida	AN11	激活	De Vetten et al., 1997
拟南芥	Arabidopsis thaliana	TTG1	激活	Walker et al., 1999
玉米	Zea mays	PAC1	激活	Carey et al., 2004
		MP1	激活	Carey et al., 2004
蒺藜苜蓿	Medicago truncatula	WD40-1	激活	Pang et al., 2009
苹果	Malus domestica	MdTTG1	激活	Brueggemann et al., 2010
石榴	Punica granatum	PgWD40	激活	Ben-Simhon et al., 2011
红薯	Ipomoea batatas	IbWD40	激活	Dong et al., 2014
葡萄	Vitis vinifera	VvWDR1/2	激活	Matus et al., 2010
马铃薯	Solanum tuberosum	StWD40	激活	Payyavula et al., 2013
		StAN11	激活	Li et al., 2014
杨梅	Myrica rubra	MrWD40-1	激活	Liu et al., 2013
紫苏	Gynura bicolor	PFWD	激活	Sompornpailin et al., 2002
甜樱桃	Prunus avium	PaWD40	激活	Starkevi? et al., 2015
圆叶牵牛	Ipomoea purpurea	WDR1	激活	Morita et al., 2006
花椰菜	Brassica oleracea var. botrytis	BoWD40	激活	Chiu and Li, 2012
苦荞麦	Fagopyrum tataricum	FtWD40	激活	Yao et al., 2017b