光照调控园艺作物花青素苷生物合成的分子机制

doi:10.16420/j.issn.0513-353x.2021-0497

摘要/Abstract

摘要：

综述了近30年来有关光强、光质和光周期等光照因素调控园艺作物花青素苷生物合成的研究进展,并侧重总结了该生物学过程中关键转录因子及其作用机制,梳理其分子调控网络。

Abstract:

Conducted a systematic review on the anthocyanin biosynthesis in horticultural crops regulated by various light factors,e.g. intensity,quality,and photoperiod,based on the literatures in the last three decades. Key transcription factors,their functional mechanism,and the entire molecular regulatory network related to this biological process were summarized.

Key words: horticultural crop, light, anthocyanin, biosynthesis

中图分类号:

洪艳, 武宇薇, 宋想, 李梦灵, 戴思兰. 光照调控园艺作物花青素苷生物合成的分子机制[J]. 园艺学报, 2021, 48(10): 1983-2000.

HONG Yan, WU Yuwei, SONG Xiang, LI Mengling, DAI Silan. Molecular Mechanism of Light-induced Anthocyanin Biosynthesis in Horticultural Crops[J]. Acta Horticulturae Sinica, 2021, 48(10): 1983-2000.

图/表 5

表1 部分园艺作物花青素苷生物合成途径响应光照表达的结构基因

Table 1 Light-induced structural genes in the anthocyanin biosynthetic pathway in horticultural crops

园艺作物 Horticultural crop	响应光诱导表达的结构基因Light-induced structural gene									参考文献 Reference
园艺作物 Horticultural crop	CHS	CHI	F3H	F3'H	F3'5'H	DFR	ANS	MT	UFGT（GT）	参考文献 Reference
荔枝 Litchi chinensis	√	√	√			√	√		√	Wei et al.,2011
葡萄 Vitis vinifera	√	√	√		√	√		√	√	Azuma et al.,2012
杨梅 Myrica rubra			√	√		√	√		√	Niu et al.,2010
番茄Solanum lycopersicum		√	√							Løvdal et al.,2010
茄子Solanum melongena	√	√	√		√	√	√			Li et al.,2018
梨 Pyrus pyrifolia						√	√			Feng et al.,2010
桃 Prunus persica						√	√			Ravaglia et al.,2013
苹果 Malus × domestica	√	√	√			√			√	Takos et al.,2006b; Feng et al.,2013
海棠 Malus sp.	√			√			√			Lu et al.,2017
百合 Lilium spp.						√				Nakatsuka et al.,2009
菊花Chrysanthemum × morifolium	√		√	√		√	√	√	√	Hong et al.,2015; Hong et al.,2016
矮牵牛 Petunia hybrida	√	√								Albert et al.,2009
叶用莴苣Lactuca sativa	√	√	√	√		√	√		√	Zhang et al.,2018b

表2 园艺作物中鉴定的响应光照表达且调控花青素苷生物合成的MYB类转录因子

Table 2 Light-induced MYB transcription factors in the anthocyanin biosynthetic pathway amongst horticultural crops

物种 Species	基因名称 Gene name	调控部位 Regulatory position	参考文献 Reference
苹果Malus × domestica	MdMYB1	果皮Skin	Takos et al.,2006a
	MdMYBA	果皮Skin	Ban et al.,2007
	MdMYB10	果肉Pulp	Feng et al.,2013
杨梅Myrica rubra	MrMYB1	果实Fruit	Niu et al.,2010
葡萄Vitis vinifera	VvMYBA1	果皮Skin	Koyama et al.,2012
葡萄Vitis vinifera	VlMYBA2	果皮Skin	Azuma et al.,2012
荔枝Litchi chinensis	LcMYB1	果皮Skin	Lai et al.,2014
桃Prunus persica	PpMYB10	果皮Skin	Ravaglia et al.,2013
梨Pyrus pyrifolia	PyMYB10	果皮Skin	Feng et al.,2010;Zhang et al.,2011
草莓Fragaria ananassa	FaMYB10	果实Fruit	Miyawaki et al.,2012
森林草莓Fragaria vesca	FvMYB10	花瓣Petal	Lin-Wang et al.,2010
番茄Solanum lycopersicum	AFT	果实Fruit	Cao et al.,2017
紫苏Perilla frutescens	PfMYB-P1	叶片Leaf	Gong et al.,1999
矮牵牛Perilla frutescens	PHZ	花瓣Petal	Albert et al.,2011
百合Lilium spp.	LrMYB15	花瓣Petal	Yamagishi,2016
菊花Chrysanthemum × morifolium	CmMYB4-7	舌状花Ray floret	Hong et al.,2016,2019

图1 光信号转导调控拟南芥花青素苷生物合成的分子机制（Laubinger et al.,2004;Stracke et al.,2010;Cloix et al.,2012;Lau & Deng,2012;Li et al.,2012,2016;Maier et al.,2013;Maier & Hoecker,2015） a:光照条件下,COP1被激活的光受体蛋白抑制,如蓝光下CRY1与SPA进行物理互作,并将SPA与COP1隔离,从而阻碍COP1-SPA蛋白复合体的形成;之后可见光促进COP1从细胞核向细胞质转运,并在很长一段时间内抑制COP1的表达,HY5不断积累并与靶蛋白结合,从而促进植物的光形态建成;b:黑暗条件下,COP1定位在细胞核内与SPA形成复合体,靶向结合bZIPHY5等,并通过蛋白酶体介导的泛素化途径使这些转录因子降解;c:无UV-B照射时,UVR8以二聚体的形式存在于细胞质和细胞核中,接收到UV-B信号后,其转移到细胞核中,蛋白构象从二聚体变为单体,并与COP1发生互作,开启UV-B信号通路,正向调节UV-B诱导类黄酮类化合物的合成。

Fig. 1 Molecular mechanism of anthocyanin biosynthesis in Arabidopsis regulated by light a:Under light condition,COP1 is suppressed by active photoreceptor proteins,e.g.,under blue light condition,CRY1 physically interacts with SAP and segregates it from COP1,which hinders the formation of COP1-SPA complex;thereafter,visible light promotes the transport of COP1 from cell nucleus to cytoplasm,which suppresses the expression of COP1 in a long time and results in the decline of COP1 activity;on the other hand,the continuous accumulation of HY5 and the binding between HY5 and targeted proteins promotes the plant photomorphosis. b:Under dark condition,COP1 integrates with SPA in cell nucleus and binds to bZIP transcription factors（e.g.,HY5）as a complex,and then degrades these transcription factors via proteasome-mediated ubiquitylation. c:Under UV-B-missing condition,UVR8 exists in cytoplasm and cell nucleus as dimer;upon receiving UV-B signal,UVR8 is transported from cytoplasm to cell nucleus,whose protein conformation is accordingly changed from dimer to monomer;thereafter,UVR8 interacts with COP1 and activites UV-B signal pathway,which positively regulates UV-B and induces the biosynthesis of flavonoids.

图2 拟南芥中光受体介导光质信号转导调控花青素苷生物合成的转导机制（Kim et al.,2003;Shin et al.,2007;Vandenbussche et al.,2007;Liu et al.,2015b）

Fig. 2 Mechanism of anthocyanin biosynthesis regulated by photoreceptor-mediated light quality signal transduction in Arabidopsis

图3 苹果中UV-B调控花青素苷生物合成的信号转导机制（Peng et al.,2013;Bai et al.,2014;An et al.,2019;Fang et al.,2019a;2019b;Hu et al.,2020）

Fig. 3 Signal transduction mechanism of anthocyanin biosynthesis regulated by UV-B in apple

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