植物类胡萝卜素代谢及调控研究进展

doi:10.16420/j.issn.0513-353x.2021-0531

摘要/Abstract

摘要：

本文概述了植物类胡萝卜素的代谢途径及关键的代谢流控制节点，重点总结了光合器官和类胡萝卜素存储器官中色素代谢的转录调控及转录后、翻译后、表观遗传等调控机制。

Abstract:

This review briefly describes the carotenoid metabolic pathway and main flux-controlling steps for carotenoid accumulation in plants. It highlights our recent understanding of the regulatory mechanisms underlying carotenoid accumulation at transcriptional，post-transcriptional，post-translational and epigenetic levels.

Key words: photosynthetic organ, storage organ, carotenoid, metabolism, regulation

中图分类号:

S6
Q 946.92

陆晨飞, 高月霞, 黄河, 戴思兰. 植物类胡萝卜素代谢及调控研究进展[J]. 园艺学报, 2022, 49(12): 2559-2578.

LU Chenfei, GAO Yuexia, HUANG He, DAI Silan. Carotenoid Metabolism and Regulation in Plants[J]. Acta Horticulturae Sinica, 2022, 49(12): 2559-2578.

图/表 3

参考文献 131

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不同类型器官 Different types of organs	信号因子 Signaling factor	物种 Species	转录因子 Transcription factor	调控机制 Regulatory mechanism	参考文献 Reference
光合器官 Photosynthetic organ	光信号 Light signaling	拟南芥 Arabidopsis thaliana	AtPIF1	直接与AtPSY启动子上的G-box元件结合从而抑制其表达 AtPIF1 repressed the transcription of AtPSY by directly binding to the G-box element	Toledo-Ortiz et al.，2010
			AtPAR1	在遮荫条件下，AtPAR1能够与AtPIF1相互作用阻止其结合至AtPSY启动子上，从而促进AtPSY的表达 AtPAR1 interacted with AtPIF1 to prevent it from binding to the AtPSY promoter，thereby promoting the expression of AtPSY	Bou-Torrent et al.，2015
			AtHY5	AtHY5能特异性识别并结合到AtPSY启动子区域的G-box元件促进其表达，在调控植物类胡萝卜素代谢过程中AtHY5与AtPIF1存在拮抗作用 AtHY5 specifically could recognize and bind to the G-box element in the AtPSY promoter to promote its expression	Toledo-Ortiz et al.，2014
	温度信号 Temperature signaling	拟南芥 Arabidopsis thaliana	AtPIF1/ AtHY5	低温条件下AtHY5蛋白含量明显增加而AtPIF1蛋白含量明显减少，使得AtPSY的表达量显著增加 The protein content of AtHY5 was significantly increased and the protein content of AtPIF1 was significantly decreased under low temperature condition，resulting in a significant increase in the expression of AtPSY	Toledo-Ortiz et al.，2014
	发育信号 Developmental cue	拟南芥 Arabidopsis thaliana	AtRAP2.2	AtRAP2.2能够特异性地结合到拟南芥AtPSY和AtPDS基因的启动子上调控其表达 AtRAP2.2 specifically bind to the promoters of AtPSY and AtPDS to regulate their expression	Welsch et al.，2007
		葡萄 Vitis vinifera	VvDRL1	在烟草中过表达VvDRL1后NtZEP1和类胡萝卜素裂解基因的表达显著减少，ABA含量下降从而使得叶片延缓衰老 Overexpression of VvDRL1 in tobacco resulted in low expression of NtZEP1 and carotenoid cleavage genes	Zhu et al.，2019
类胡萝卜素存储器官 Carotenoid storage organ	光信号 Light signaling	番茄 Solanum lycopersicum	SlDET1/ SlDDB1	通过RNAi抑制SlDET1和SlDDB1表达后成熟番茄果实中的类胡萝卜素（β-胡萝卜素、番茄红素）和类黄酮含量显著上升 The contents of carotenoids and flavonoids in tomato were significantly increased after inhibiting the expression of SlDET1 and SlDDB1 by RNAi	Davuluri et al.，2005;Bino et al.，2005; Calvenzani et al.，2010
			SlBBX20	SlBBX20通过直接调节SlPSY1的表达从而影响色素生成 SlBBX20 affected carotenoid metabolism by directly regulating the expression of SlPSY1	Xiong et al.，2019
			SlPIF1a	SlPIF1a可直接与SlPSY1基因启动子上的PBE-box结合以抑制其表达 SlPIF1a could directly bind to the PBE-box in the SlPSY1 promoter to inhibit its expression	Llorente et al.， 2016
			SlPRE2	SlPRE2的表达受强光照条件抑制，当SlPRE2在番茄中过表达时，叶绿素以及类胡萝卜素合成基因SlPSY1、SlPDS、SlZDS的表达量显著下降 Overexpression of SlPRE2 resulted in low expression levels of chlorophyll and carotenoid metabolic genes SlPSY1，SlPDS and SlZDS	Zhu et al.，2017
		木瓜 Carica papaya	CpbHLH1/ CpbHLH2	CpbHLH1、CpbHLH2分别作为转录抑制子和转录激活子起始下游结构基因CpCYCB和CpLCYB的转录，且两者受到光信号的调控CpbHLH1 and CpbHLH2，as transcriptional repressors and transcriptional activators respectively，initiated the transcription of downstream structural genes CpCYCB and CpLCYB	Zhou et al.，2019
		胡萝卜 Daucus carota subsp. carota	DcPEL	QTL定位及GWAS分析显示光形态建成抑制因子PEL与胡萝卜肉质根中的类胡萝卜素积累密切相关 QTL mapping and GWAS analysis revealed that the photomorphogenesis inhibitor PEL was closely related to carotenoid accumulation in carrot	Iorizzo et al.，2016
	发育信号 Developmental cue	番茄 Solanum lycopersicum	RIN/TAGL1/ FUL1/FUL2	这些转录因子可相互作用形成不同的蛋白复合体，同时都可直接或间接的激活SlPSY1、SlPSY2、SlZDS、SlZ-ISO、SlCRTISO和SlBCH基因的转录，抑制S1LCYB、SlLCYE和S1CYCB的表达 These transcription factors have both overlapping and individual contributions to the expression of carotenoid metabolic genes，with the total effect being the positive regulation of SlPSY1，SlPSY2，SlZDS，SlZ-ISO，SlCRTISO，and SlBCH，and the negative regulation of SlLCYB，SlLCYE，and SlCYCB	Itkin et al.，2009;Martel et al.，2011; Shima et al.，2013;Fujisawa et al.，2014;Stanley & Yuan，2019
		柑橘 Citrus sinensis	CsMADS5/ CsMADS6	CsMADS5和CsMADS6作为转录激活因子通过直接结合至下游结构基因的启动子上激活其表达从而促进柑橘果实类胡萝卜素的积累 As the positive regulators，CsMADS5 and CsMADS6 promoted the accumulation of carotenoids in citrus by directly binding to the promoters of downstream structural genes	Lu et al.，2018，2021
		青瓯柑 Citrus reticulata	CrMYB68	CrMYB68可直接抑制CrBCH2和CrNCED5的表达CrMYB68 inhibited the expression of CrBCH2 and CrNCED5 directly	Zhu et al.，2017
		木瓜 Carica papaya	CpSBP1	CpSBP1可结合到CpPDS的启动子区域并激活其表达CpSBP1 could bind to the promoter region of CpPDS to activate its expression	Han et al.，2019
		猕猴桃 Actinidia deliciosa	AdMYB7	AdMYB7是直接调节AdLCYB转录的重要调控因子，当AdMYB7在烟草中过表达时，类胡萝卜素、叶绿素总含量明显增加 AdMYB7 was an important regulator that directly regulated the transcription of AdLCYB	Ampomah- Dwamena et al.，2019
		苜蓿 Medicago truncatula	MtWP1	MtWP1可直接调控花青素、类胡萝卜素合成基因如MtLCYB、MtLCYE的表达，通过影响两类色素的积累而决定花瓣呈色 MtWP1 could directly regulate the expression of anthocyanin and carotenoid metabolic genes such as MtLCYB and MtLCYE	Meng et al.，2019
		烟草 Nicotiana tabacum	NtMYB305	NtMYB305与蜜腺中的β-类胡萝卜素积累密切相关NtMYB305 was closely involved in β-carotene accumulation in nectaries	Liu et al.，2009;Wang et al，2014
		猴面花 Mimulus lewisii	MlRCP1/ MlRCP2	利用RNAi分别抑制MlRCP1和MlRCP2表达后，花瓣中所有与类胡萝卜素合成相关基因的表达量都下调，类胡萝卜素的总含量降低 The expression levels of all carotenoid metabolic genes in petals were down-regulated after inhibiting the expression of MlRCP1 and MlRCP2 by RNAi	Sagawa et al.，2016;Stanley et al.，2020
	激素信号 Phytohormones	番茄 Solanum lycopersicum	SlAP2a	SlAP2a能够激活类胡萝卜素代谢基因SlPSY1、SlCRTISO、SlBCH和SlPDS1的表达，抑制S1ZEP1和S1CYCB的转录 SlAP2a could activate the expression of carotenoid metabolic genes SlPSY1，SlCRTISO，SlBCH and SlPDS1，and inhibit the transcription of S1ZEP1 and S1CYCB	Chung et al.，2010;Karlova et al.，2011
			SIERF6	利用RNAi抑制SIERF6的表达，总类胡萝卜素及乙烯含量均显著上升 The content of total carotenoids and ethylene was significantly increased after inhibiting the expression of SIERF6 by RNAi	Lee et al.，2012
			SlNAC1/ SlNAC4	SlNAC1可直接与SlPSY1和乙烯生物合成基因的启动子相结合起抑制转录作用;SlNAC4可促进SlPSY1的表达，利用RNAi抑制SlNAC4的转录后，SlPSY1的表达量显著降低 SlNAC1 could directly bind to the promoters of SlPSY1 and ethylene biosynthesis genes to inhibit their transcription; SlNAC4 could promote the expression of SlPSY1	Ma et al.，2014;Zhu et al.，2014;Meng et al.，2016
			SlARF2A/ SlARF2B	利用RNAi抑制SlARF2A、SlARF2B的转录后，SlRIN、SlTAGL1、SlFUL1/2等参与果实成熟的发育因子表达量下降，同时类胡萝卜素代谢相关基因的表达也受到影响 The expression levels of developmental genes such as SlRIN，SlTAGL1，SlFUL1/2 and the carotenoid metabolic genes were affected after inhibiting the transcription of SlARF2A and SlARF2B by RNAi	Hao et al.，2015
			SlIPT4	SlIPT4可促进番茄SlZ-ISO和SlZDS的表达从而诱导番茄红素的积累 SlIPT4 promoted the expression of tomato SlZ-ISO and SlZDS to induce lycopene accumulation	Zhang et al.，2018
			SlBZR1	在番茄中过表达油菜素内酯响应因子SlBZR1，SlPSY1和SlZDS的转录水平上调，类胡萝卜素、可溶性糖和抗坏血酸大量积累 Overexpression of SlBZR1 in tomato resulted in up-regulated transcription of SlPSY1 and SlZDS	Liu et al.，2014
		柑橘 Citrus sinensis	CsERF61	CsERF61的表达受乙烯的诱导，该基因所编码的转录因子可直接结合至CsLCYB2的启动子上激活其表达 CsERF61 can directly bind to the promoter of CsLCYB2 to activate its expression	Zhu et al.，2021
		木瓜 Carica papaya	CpNAC1/ CpNAC2	CpNAC2可与乙烯响应蛋白CpEIN3a相互作用共同促进下游结构基因CpPDS2/4、CpLCYE、CpCHYB的表达 CpNAC2 could interact with ethylene-responsive protein CpEIN3a to promote the expression of downstream structural genes CpPDS2/4，CpLCYE and CpCHYB	Fu et al.，2016，2017
		温州蜜柑 Citrus unshiu	CubHLH1	在番茄中过表达BR信号通路负调节因子CubHLH1后类胡萝卜素生物合成基因的表达受到抑制，番茄红素含量显著降低 Overexpression of CubHLH1 in tomato inhibited the expression of carotenoid biosynthesis genes	Endo et al.，2016

不同类型器官 Different types of organs	信号因子 Signaling factor	物种 Species	转录因子 Transcription factor	调控机制 Regulatory mechanism	参考文献 Reference
光合器官 Photosynthetic organ	光信号 Light signaling	拟南芥 Arabidopsis thaliana	AtPIF1	直接与AtPSY启动子上的G-box元件结合从而抑制其表达 AtPIF1 repressed the transcription of AtPSY by directly binding to the G-box element	Toledo-Ortiz et al.，2010
			AtPAR1	在遮荫条件下，AtPAR1能够与AtPIF1相互作用阻止其结合至AtPSY启动子上，从而促进AtPSY的表达 AtPAR1 interacted with AtPIF1 to prevent it from binding to the AtPSY promoter，thereby promoting the expression of AtPSY	Bou-Torrent et al.，2015
			AtHY5	AtHY5能特异性识别并结合到AtPSY启动子区域的G-box元件促进其表达，在调控植物类胡萝卜素代谢过程中AtHY5与AtPIF1存在拮抗作用 AtHY5 specifically could recognize and bind to the G-box element in the AtPSY promoter to promote its expression	Toledo-Ortiz et al.，2014
	温度信号 Temperature signaling	拟南芥 Arabidopsis thaliana	AtPIF1/ AtHY5	低温条件下AtHY5蛋白含量明显增加而AtPIF1蛋白含量明显减少，使得AtPSY的表达量显著增加 The protein content of AtHY5 was significantly increased and the protein content of AtPIF1 was significantly decreased under low temperature condition，resulting in a significant increase in the expression of AtPSY	Toledo-Ortiz et al.，2014
	发育信号 Developmental cue	拟南芥 Arabidopsis thaliana	AtRAP2.2	AtRAP2.2能够特异性地结合到拟南芥AtPSY和AtPDS基因的启动子上调控其表达 AtRAP2.2 specifically bind to the promoters of AtPSY and AtPDS to regulate their expression	Welsch et al.，2007
		葡萄 Vitis vinifera	VvDRL1	在烟草中过表达VvDRL1后NtZEP1和类胡萝卜素裂解基因的表达显著减少，ABA含量下降从而使得叶片延缓衰老 Overexpression of VvDRL1 in tobacco resulted in low expression of NtZEP1 and carotenoid cleavage genes	Zhu et al.，2019
类胡萝卜素存储器官 Carotenoid storage organ	光信号 Light signaling	番茄 Solanum lycopersicum	SlDET1/ SlDDB1	通过RNAi抑制SlDET1和SlDDB1表达后成熟番茄果实中的类胡萝卜素（β-胡萝卜素、番茄红素）和类黄酮含量显著上升 The contents of carotenoids and flavonoids in tomato were significantly increased after inhibiting the expression of SlDET1 and SlDDB1 by RNAi	Davuluri et al.，2005;Bino et al.，2005; Calvenzani et al.，2010
			SlBBX20	SlBBX20通过直接调节SlPSY1的表达从而影响色素生成 SlBBX20 affected carotenoid metabolism by directly regulating the expression of SlPSY1	Xiong et al.，2019
			SlPIF1a	SlPIF1a可直接与SlPSY1基因启动子上的PBE-box结合以抑制其表达 SlPIF1a could directly bind to the PBE-box in the SlPSY1 promoter to inhibit its expression	Llorente et al.， 2016
			SlPRE2	SlPRE2的表达受强光照条件抑制，当SlPRE2在番茄中过表达时，叶绿素以及类胡萝卜素合成基因SlPSY1、SlPDS、SlZDS的表达量显著下降 Overexpression of SlPRE2 resulted in low expression levels of chlorophyll and carotenoid metabolic genes SlPSY1，SlPDS and SlZDS	Zhu et al.，2017
		木瓜 Carica papaya	CpbHLH1/ CpbHLH2	CpbHLH1、CpbHLH2分别作为转录抑制子和转录激活子起始下游结构基因CpCYCB和CpLCYB的转录，且两者受到光信号的调控CpbHLH1 and CpbHLH2，as transcriptional repressors and transcriptional activators respectively，initiated the transcription of downstream structural genes CpCYCB and CpLCYB	Zhou et al.，2019
		胡萝卜 Daucus carota subsp. carota	DcPEL	QTL定位及GWAS分析显示光形态建成抑制因子PEL与胡萝卜肉质根中的类胡萝卜素积累密切相关 QTL mapping and GWAS analysis revealed that the photomorphogenesis inhibitor PEL was closely related to carotenoid accumulation in carrot	Iorizzo et al.，2016
	发育信号 Developmental cue	番茄 Solanum lycopersicum	RIN/TAGL1/ FUL1/FUL2	这些转录因子可相互作用形成不同的蛋白复合体，同时都可直接或间接的激活SlPSY1、SlPSY2、SlZDS、SlZ-ISO、SlCRTISO和SlBCH基因的转录，抑制S1LCYB、SlLCYE和S1CYCB的表达 These transcription factors have both overlapping and individual contributions to the expression of carotenoid metabolic genes，with the total effect being the positive regulation of SlPSY1，SlPSY2，SlZDS，SlZ-ISO，SlCRTISO，and SlBCH，and the negative regulation of SlLCYB，SlLCYE，and SlCYCB	Itkin et al.，2009;Martel et al.，2011; Shima et al.，2013;Fujisawa et al.，2014;Stanley & Yuan，2019
		柑橘 Citrus sinensis	CsMADS5/ CsMADS6	CsMADS5和CsMADS6作为转录激活因子通过直接结合至下游结构基因的启动子上激活其表达从而促进柑橘果实类胡萝卜素的积累 As the positive regulators，CsMADS5 and CsMADS6 promoted the accumulation of carotenoids in citrus by directly binding to the promoters of downstream structural genes	Lu et al.，2018，2021
		青瓯柑 Citrus reticulata	CrMYB68	CrMYB68可直接抑制CrBCH2和CrNCED5的表达CrMYB68 inhibited the expression of CrBCH2 and CrNCED5 directly	Zhu et al.，2017
		木瓜 Carica papaya	CpSBP1	CpSBP1可结合到CpPDS的启动子区域并激活其表达CpSBP1 could bind to the promoter region of CpPDS to activate its expression	Han et al.，2019
		猕猴桃 Actinidia deliciosa	AdMYB7	AdMYB7是直接调节AdLCYB转录的重要调控因子，当AdMYB7在烟草中过表达时，类胡萝卜素、叶绿素总含量明显增加 AdMYB7 was an important regulator that directly regulated the transcription of AdLCYB	Ampomah- Dwamena et al.，2019
		苜蓿 Medicago truncatula	MtWP1	MtWP1可直接调控花青素、类胡萝卜素合成基因如MtLCYB、MtLCYE的表达，通过影响两类色素的积累而决定花瓣呈色 MtWP1 could directly regulate the expression of anthocyanin and carotenoid metabolic genes such as MtLCYB and MtLCYE	Meng et al.，2019
		烟草 Nicotiana tabacum	NtMYB305	NtMYB305与蜜腺中的β-类胡萝卜素积累密切相关NtMYB305 was closely involved in β-carotene accumulation in nectaries	Liu et al.，2009;Wang et al，2014
		猴面花 Mimulus lewisii	MlRCP1/ MlRCP2	利用RNAi分别抑制MlRCP1和MlRCP2表达后，花瓣中所有与类胡萝卜素合成相关基因的表达量都下调，类胡萝卜素的总含量降低 The expression levels of all carotenoid metabolic genes in petals were down-regulated after inhibiting the expression of MlRCP1 and MlRCP2 by RNAi	Sagawa et al.，2016;Stanley et al.，2020
	激素信号 Phytohormones	番茄 Solanum lycopersicum	SlAP2a	SlAP2a能够激活类胡萝卜素代谢基因SlPSY1、SlCRTISO、SlBCH和SlPDS1的表达，抑制S1ZEP1和S1CYCB的转录 SlAP2a could activate the expression of carotenoid metabolic genes SlPSY1，SlCRTISO，SlBCH and SlPDS1，and inhibit the transcription of S1ZEP1 and S1CYCB	Chung et al.，2010;Karlova et al.，2011
			SIERF6	利用RNAi抑制SIERF6的表达，总类胡萝卜素及乙烯含量均显著上升 The content of total carotenoids and ethylene was significantly increased after inhibiting the expression of SIERF6 by RNAi	Lee et al.，2012
			SlNAC1/ SlNAC4	SlNAC1可直接与SlPSY1和乙烯生物合成基因的启动子相结合起抑制转录作用;SlNAC4可促进SlPSY1的表达，利用RNAi抑制SlNAC4的转录后，SlPSY1的表达量显著降低 SlNAC1 could directly bind to the promoters of SlPSY1 and ethylene biosynthesis genes to inhibit their transcription; SlNAC4 could promote the expression of SlPSY1	Ma et al.，2014;Zhu et al.，2014;Meng et al.，2016
			SlARF2A/ SlARF2B	利用RNAi抑制SlARF2A、SlARF2B的转录后，SlRIN、SlTAGL1、SlFUL1/2等参与果实成熟的发育因子表达量下降，同时类胡萝卜素代谢相关基因的表达也受到影响 The expression levels of developmental genes such as SlRIN，SlTAGL1，SlFUL1/2 and the carotenoid metabolic genes were affected after inhibiting the transcription of SlARF2A and SlARF2B by RNAi	Hao et al.，2015
			SlIPT4	SlIPT4可促进番茄SlZ-ISO和SlZDS的表达从而诱导番茄红素的积累 SlIPT4 promoted the expression of tomato SlZ-ISO and SlZDS to induce lycopene accumulation	Zhang et al.，2018
			SlBZR1	在番茄中过表达油菜素内酯响应因子SlBZR1，SlPSY1和SlZDS的转录水平上调，类胡萝卜素、可溶性糖和抗坏血酸大量积累 Overexpression of SlBZR1 in tomato resulted in up-regulated transcription of SlPSY1 and SlZDS	Liu et al.，2014
		柑橘 Citrus sinensis	CsERF61	CsERF61的表达受乙烯的诱导，该基因所编码的转录因子可直接结合至CsLCYB2的启动子上激活其表达 CsERF61 can directly bind to the promoter of CsLCYB2 to activate its expression	Zhu et al.，2021
		木瓜 Carica papaya	CpNAC1/ CpNAC2	CpNAC2可与乙烯响应蛋白CpEIN3a相互作用共同促进下游结构基因CpPDS2/4、CpLCYE、CpCHYB的表达 CpNAC2 could interact with ethylene-responsive protein CpEIN3a to promote the expression of downstream structural genes CpPDS2/4，CpLCYE and CpCHYB	Fu et al.，2016，2017
		温州蜜柑 Citrus unshiu	CubHLH1	在番茄中过表达BR信号通路负调节因子CubHLH1后类胡萝卜素生物合成基因的表达受到抑制，番茄红素含量显著降低 Overexpression of CubHLH1 in tomato inhibited the expression of carotenoid biosynthesis genes	Endo et al.，2016