葡萄类黄酮代谢研究进展

doi:10.16420/j.issn.0513-353x.2021-0558

摘要/Abstract

摘要：

类黄酮是葡萄中含量最丰富的一类次生代谢物,在植株生长发育、抗紫外线和病虫害、果实色泽、风味品质以及营养价值等方面发挥着至关重要的作用。葡萄果实中类黄酮的种类多样,积累特点具有品种、组织和发育特异性。葡萄类黄酮代谢主要受UDP-葡萄糖：类黄酮-3-O-葡萄糖基转移酶（UFGT）和MYBA1等基因的表达调控,还受到多种生物和非生物因素的影响。从生物学功能、种类和积累特点、合成与代谢调控以及影响因素这几个方面综述了葡萄类黄酮代谢研究的进展,从完善类黄酮含量数据库、揭示重要基因功能、解析代谢调控网络、探索多因素交互作用机理这几个方面对今后葡萄类黄酮代谢研究方向提出了展望。

关键词: 葡萄, 类黄酮, 次生代谢, 果实品质

Abstract:

Flavonoids are the most abundant secondary metabolites in grape. These compounds play vital roles in grape growth and development,resistance to ultraviolet radiation and diseases/insects,berry coloration and flavor quality,as well as nutritional values. Grape berries possess various types of flavonoids,which show variety-,tissue-,and development-specific accumulation characteristics. Flavonoid metabolism in grape is mainly regulated by UDP-glucose：flavonoid 3-O-glycosyltransferase（UFGT）and MYBA1 genes,and also affected by a variety of biotic and abiotic factors. The review summarizes recent data related to the flavonoid metabolism from the aspects of biological functions,composition and accumulation characteristics,biosynthesis and regulation mechanisms,and influencing factors. Finally,the future research directions of grape flavonoid metabolism are also put forward from the aspects of perfecting flavonoid content database,uncovering important gene functions,analyzing metabolic regulation networks,and exploring the interaction mechanisms of multiple factors.

Key words: grape, flavonoids, secondary metabolism, fruit quality

中图分类号:

S663.1

卢素文, 郑暄昂, 王佳洋, 房经贵. 葡萄类黄酮代谢研究进展[J]. 园艺学报, 2021, 48(12): 2506-2524.

LU Suwen, ZHENG Xuanang, WANG Jiayang, FANG Jinggui. Research Progress on the Metabolism of Flavonoids in Grape[J]. Acta Horticulturae Sinica, 2021, 48(12): 2506-2524.

图/表 4

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物质类别 Class	结构单体 Monomer	衍生物 Derivative
花青苷 Anthocyanin	飞燕草色素Delphinidin（Dp）	3-O-葡萄糖苷3-O-glucoside^*
	矢车菊色素Cyanidin（Cy）	3-O-（6-O-乙酰）-葡萄糖苷3-O-(6-O-acetyl)-glucoside
	牵牛花色素Petunidin（Pt）芍药色素Peonidin（Pn）锦葵色素Malvidin（Mv）^*	3-O-（6-O-香豆酰）-葡萄糖苷3-O-(6-O-coumaroyl)-glucoside 3-（6-O-咖啡酰）-葡萄糖苷3-O-(6-O-caffeoyl)-glucoside
黄酮醇 Flavonol	山奈酚Kaempferol	3-O-葡萄糖苷3-O-glucoside^*
	槲皮素Quercetin ^*	3-O-葡萄糖苷酸3-O-glucuronide
	杨梅素Myricetin 异鼠李素Isorhamnetin 落叶松素Laricitrin 丁香亭Syringetin	3-O-半乳糖苷3-O-galactoside 3-O-芸香糖苷3-O-rutinoside 3-O-鼠李糖苷3-O-rhamnoside 3-（6-O-乙酰）葡萄糖苷3-(6-O-acetyl)glucoside 3-（6-O-香豆酰）葡萄糖苷3-(6-O-coumaroyl)glucoside
黄烷醇 Flavanol	儿茶素Catechin（C）^* 表儿茶素Epicatechin（EC）表儿茶素没食子酸酯Epicatechin gallate（ECG）	原花青素B1 Procyanidin B1 ^* 原花青素B2 Procyanidin B2

物质类别 Class	结构单体 Monomer	衍生物 Derivative
花青苷 Anthocyanin	飞燕草色素Delphinidin（Dp）	3-O-葡萄糖苷3-O-glucoside^*
	矢车菊色素Cyanidin（Cy）	3-O-（6-O-乙酰）-葡萄糖苷3-O-(6-O-acetyl)-glucoside
	牵牛花色素Petunidin（Pt）芍药色素Peonidin（Pn）锦葵色素Malvidin（Mv）^*	3-O-（6-O-香豆酰）-葡萄糖苷3-O-(6-O-coumaroyl)-glucoside 3-（6-O-咖啡酰）-葡萄糖苷3-O-(6-O-caffeoyl)-glucoside
黄酮醇 Flavonol	山奈酚Kaempferol	3-O-葡萄糖苷3-O-glucoside^*
	槲皮素Quercetin ^*	3-O-葡萄糖苷酸3-O-glucuronide
	杨梅素Myricetin 异鼠李素Isorhamnetin 落叶松素Laricitrin 丁香亭Syringetin	3-O-半乳糖苷3-O-galactoside 3-O-芸香糖苷3-O-rutinoside 3-O-鼠李糖苷3-O-rhamnoside 3-（6-O-乙酰）葡萄糖苷3-(6-O-acetyl)glucoside 3-（6-O-香豆酰）葡萄糖苷3-(6-O-coumaroyl)glucoside
黄烷醇 Flavanol	儿茶素Catechin（C）^* 表儿茶素Epicatechin（EC）表儿茶素没食子酸酯Epicatechin gallate（ECG）	原花青素B1 Procyanidin B1 ^* 原花青素B2 Procyanidin B2

基因 Gene	基因编号 Gene ID	基因描述 Description	碱基/bp Nucleotide base		氨基酸 Amino acid	分子量/ kD Molecular weight	等电点 Isoelectric point	外显子 Exon	保守结构域/Pfam Conserved domains
PAL	VIT_06s0004g02620	苯丙氨酸解氨酶 Phenylalanine ammonia lyase	2 421		717	77.97	6.12	2	PF00221 芳香族氨基酸裂解酶 Aromatic amino acid lyase
C4H	VIT_06s0004g08150	肉桂酸4-羟化酶 Cinnamate4-hydroxylase	1 695		505	57.69	9.43	3	PF00067 细胞色素P450 Cytochrome P450
4CL	VIT_16s0039g02040	4-香豆酰CoA 连接酶 4-coumarate:CoA ligase	1 980		548	59.49	5.77	6	PF00501 AMP依赖的合成酶/连接酶 AMP-dependent synthetase/ ligase
CHS	VIT_05s0136g00260	查尔酮合成酶 Chalcone synthase	1 369		389	42.60	6.58	2	PF00195 查耳酮/芪合酶 Chalcone/stilbene synthase
CHI	VIT_13s0067g02870	查尔酮异构酶 Chalcone flavanone isomerase	1 081		247	27.47	4.58	5	PF02431 查尔酮异构酶 Chalcone isomerase
F3H	VIT_04s0023g03370	黄烷酮3羟化酶Flavanone 3-hydroxylase	1 319		363	40.81	5.21	3	PF14226 非血红素双加氧酶 Non-haem dioxygenase
F3'5'H	VIT_06s0009g02840	黄烷酮3ʹ 5ʹ羟化酶 Flavonoid 3ʹ,5ʹ-hydroxylase	1 826		508	57.05	8.46	2	PF00067 细胞色素P450 Cytochrome P450
基因 Gene	基因编号 Gene ID	基因描述 Description		碱基/bp Nucleotide base	氨基酸 Amino acid	分子量/ kD Molecular weight	等电点 Isoelectric point	外显子 Exon	保守结构域/Pfam Conserved domains
F3'H	VIT_17s0000g07200	黄烷酮3ʹ羟化酶 Flavonoid 3ʹ-hydroxylase		1 774	451	50.28	8.14	4	PF00067 细胞色素P450 Cytochrome P450
DFR	VIT_18s0001g12800	二氢黄酮醇4-还原酶Dihydroflavonol 4-reductase		1 435	375	41.94	6.75	6	PF01370 NAD依赖的差向异构酶/脱水酶NAD-dependent epimerase / dehydratase
LAR	VIT_17s0000g04150	无色花色素还原酶 Leucoanthocyanidin reductase		1 364	358	38.70	4.83	5	PF05368 NmrA类似结构域 NmrA-like domain
ANR	VIT_02s0025g01260	花色素还原酶 Anthocyanidin reductase		1 334	342	37.88	5.01	5	PF01370 NAD依赖的差向异构酶/脱水酶NAD-dependent epimerase / dehydratase
LDOX	VIT_02s0025g04720	无色花色素双加氧酶 Leucoanthocyanidin dioxgenase		1 298	355	40.19	5.69	2	PF14226 非血红素双加氧酶 Non-haem dioxygenase
FLS	VIT_18s0001g03490	黄酮醇合酶 Flavonol synthase		1 015	335	38.39	6.66	3	PF14226 非血红素双加氧酶N端结构域Non-haem dioxygenase N-terminal domain
UFGT	VIT_16s0039g02230	UDP-葡萄糖：类黄酮-3-O-葡萄糖基转移酶 UDP-glucose：flavonoid 3-O-glucosyltransferase		1 435	456	50.03	6.19	2	PF00201 UDP-葡萄糖醛酸基/葡糖基转移酶UDP-glucuronosyl / UDP-glucosyltransferase
OMT	VIT_01s0010g03510	O-甲基转移酶 O-methyltransferase		876	235	26.50	7.13	5	PF01596 I类硫腺苷甲硫氨酸依赖的 O-甲基转移酶 Class I-like SAM-dependent O-methyltransferase
AT	VIT_03s0017g00870	花色苷酰基转移酶 Anthocyanin-glucoside-acyltransferase		1 284	427	47.50	7.65	1	PF02458 转移酶Transferase
GST	VIT_04s0079g00690	谷胱甘肽-S-转移酶 Glutathione S-transferase		831	213	24.22	5.69	3	PF00043 谷胱甘肽转移酶 Glutathione S-transferase
MATE	VIT_16s0050g00900	MATE-type转运子 AnthoMATE（AM）transporter		1 589	489	53.49	6.12	7	PF01554 多药和毒性化合物外排转运蛋白Multi antimicrobial extrusion protein
ABC	VIT_16s0050g02480	ABC转运蛋白 ABC transporter		4 990	1 480	164.81	6.80	12	PF00664 ABC转运子1型 ABC transporter type 1

基因 Gene	基因编号 Gene ID	基因描述 Description	碱基/bp Nucleotide base		氨基酸 Amino acid	分子量/ kD Molecular weight	等电点 Isoelectric point	外显子 Exon	保守结构域/Pfam Conserved domains
PAL	VIT_06s0004g02620	苯丙氨酸解氨酶 Phenylalanine ammonia lyase	2 421		717	77.97	6.12	2	PF00221 芳香族氨基酸裂解酶 Aromatic amino acid lyase
C4H	VIT_06s0004g08150	肉桂酸4-羟化酶 Cinnamate4-hydroxylase	1 695		505	57.69	9.43	3	PF00067 细胞色素P450 Cytochrome P450
4CL	VIT_16s0039g02040	4-香豆酰CoA 连接酶 4-coumarate:CoA ligase	1 980		548	59.49	5.77	6	PF00501 AMP依赖的合成酶/连接酶 AMP-dependent synthetase/ ligase
CHS	VIT_05s0136g00260	查尔酮合成酶 Chalcone synthase	1 369		389	42.60	6.58	2	PF00195 查耳酮/芪合酶 Chalcone/stilbene synthase
CHI	VIT_13s0067g02870	查尔酮异构酶 Chalcone flavanone isomerase	1 081		247	27.47	4.58	5	PF02431 查尔酮异构酶 Chalcone isomerase
F3H	VIT_04s0023g03370	黄烷酮3羟化酶Flavanone 3-hydroxylase	1 319		363	40.81	5.21	3	PF14226 非血红素双加氧酶 Non-haem dioxygenase
F3'5'H	VIT_06s0009g02840	黄烷酮3ʹ 5ʹ羟化酶 Flavonoid 3ʹ,5ʹ-hydroxylase	1 826		508	57.05	8.46	2	PF00067 细胞色素P450 Cytochrome P450
基因 Gene	基因编号 Gene ID	基因描述 Description		碱基/bp Nucleotide base	氨基酸 Amino acid	分子量/ kD Molecular weight	等电点 Isoelectric point	外显子 Exon	保守结构域/Pfam Conserved domains
F3'H	VIT_17s0000g07200	黄烷酮3ʹ羟化酶 Flavonoid 3ʹ-hydroxylase		1 774	451	50.28	8.14	4	PF00067 细胞色素P450 Cytochrome P450
DFR	VIT_18s0001g12800	二氢黄酮醇4-还原酶Dihydroflavonol 4-reductase		1 435	375	41.94	6.75	6	PF01370 NAD依赖的差向异构酶/脱水酶NAD-dependent epimerase / dehydratase
LAR	VIT_17s0000g04150	无色花色素还原酶 Leucoanthocyanidin reductase		1 364	358	38.70	4.83	5	PF05368 NmrA类似结构域 NmrA-like domain
ANR	VIT_02s0025g01260	花色素还原酶 Anthocyanidin reductase		1 334	342	37.88	5.01	5	PF01370 NAD依赖的差向异构酶/脱水酶NAD-dependent epimerase / dehydratase
LDOX	VIT_02s0025g04720	无色花色素双加氧酶 Leucoanthocyanidin dioxgenase		1 298	355	40.19	5.69	2	PF14226 非血红素双加氧酶 Non-haem dioxygenase
FLS	VIT_18s0001g03490	黄酮醇合酶 Flavonol synthase		1 015	335	38.39	6.66	3	PF14226 非血红素双加氧酶N端结构域Non-haem dioxygenase N-terminal domain
UFGT	VIT_16s0039g02230	UDP-葡萄糖：类黄酮-3-O-葡萄糖基转移酶 UDP-glucose：flavonoid 3-O-glucosyltransferase		1 435	456	50.03	6.19	2	PF00201 UDP-葡萄糖醛酸基/葡糖基转移酶UDP-glucuronosyl / UDP-glucosyltransferase
OMT	VIT_01s0010g03510	O-甲基转移酶 O-methyltransferase		876	235	26.50	7.13	5	PF01596 I类硫腺苷甲硫氨酸依赖的 O-甲基转移酶 Class I-like SAM-dependent O-methyltransferase
AT	VIT_03s0017g00870	花色苷酰基转移酶 Anthocyanin-glucoside-acyltransferase		1 284	427	47.50	7.65	1	PF02458 转移酶Transferase
GST	VIT_04s0079g00690	谷胱甘肽-S-转移酶 Glutathione S-transferase		831	213	24.22	5.69	3	PF00043 谷胱甘肽转移酶 Glutathione S-transferase
MATE	VIT_16s0050g00900	MATE-type转运子 AnthoMATE（AM）transporter		1 589	489	53.49	6.12	7	PF01554 多药和毒性化合物外排转运蛋白Multi antimicrobial extrusion protein
ABC	VIT_16s0050g02480	ABC转运蛋白 ABC transporter		4 990	1 480	164.81	6.80	12	PF00664 ABC转运子1型 ABC transporter type 1

转录因子家族 TF family	基因 Gene	基因编号 Gene ID	蛋白编号 Protein ID	Ensemble编号 Ensemble ID	调控方式 Regulative way	参考文献 Reference
MYB	MYBA1	AB097923	BAD18977	VIT_ 02s0033g00410	激活UFGT、CHS3、LDOX、3AT,促进花青苷合成Activates the expression of UFGT,CHS3,LDOX and 3AT,to promote anthocyanin synthesis	Kobayashi et al.,2004;Rinaldo et al.,2015;Poudel et al.,2021
	MYBA2	AB097924	BAD18978	VIT_ 02s0033g00390	激活UFGT、CHS3、LDOX,促进花青苷合成Activates the expression of UFGT,CHS3,LDOX and 3AT,to promote anthocyanin synthesis	Kobayashi et al.,2004;Poudel et al.,2021
	MYBA3	AB097925	BAD18979	VIT_ 02s0033g00450	未知Unknown	Kobayashi et al.,2004
	MYBPA1	AM259485	CAJ90831	VIT_ 15s0046g00170	激活ANR、LAR1、F3'5'H、LDOX、CHI,不激活UFGT,促进种子和果实发育早期原花青素合成Activates the expression of ANR, LAR1,F3'5'H,LDOX and CHI,but not UFGT,to promote proanthocyanidin synthesis in seed and early fruit development	Bogs et al.,2007;Terrier et al.,2009
	MYBPA2	EU919682	ACK56131	VIT_ 11s0016g01310	激活ANR、LAR1,促进果实原花青素合成Activates the expression of ANR and LAR1,to promote proanthocyanidin synthesis in fruit	Terrier et al.,2009
	MYBPAR	AB911341	BAP39802	VIT_ 11s0016g01300	激活ANR、LAR1、LAR2、CHS3、F3'5'H、GT1、SCP、MATE,不激活UFGT,促进果实原花青素合成Activates the expression of ANR,LAR1,LAR2,CHS3,F3'5'H,GT1,SCP and MATE,but not UFGT,to promote proanthocyanidin synthesis in fruit	Koyama et al.,2014
	MYB5a	AY555190	AAS68190	VIT_ 08s0007g07230	与AtEGL3（bHLH蛋白）互作,激活LAR1、CHI、F3'5'H、ANS,促进花青苷、黄酮醇和原花青素合成Interacts with AtEGL3（a bHLH protein）and activates the expression of LAR1,CHI,F3'5'Hand ANS,to promote the synthesis of anthocyanin,flavonol,and procyanidin	Deluc et al.,2006,2008;
转录因子家族 TF family	基因 Gene	基因编号 Gene ID	蛋白编号 Protein ID	Ensemble编号 Ensemble ID	调控方式 Regulative way	参考文献 Reference
	MYB5b	AY899404	AAX51291	VIT_ 06s0004g00570	与AtEGL3（bHLH蛋白）互作,激活ANR、LAR1、CHI、F3'5'H、ANS,促进花青苷、黄酮醇和原花青素合成Interacts with AtEGL3（a bHLH protein）and activates the expression of ANR,LAR1,CHI,F3'5'H and ANS,to promote the synthesis of anthocyanin,flavonol,and procyanidin	Deluc et al.,2008
	MYBF1	GQ423422	ACV81697	VIT_ 07s0005g01210	激活CHS、CHI、FLS,促进黄酮醇合成Activates the expression of CHS,CHI and FLS,to promote flavonol synthesis	Czemmel et al. 2009
	MYB4	EF113078	ABL61515	VIT_ 01s0011g04760	抑制花青素合成 Inhibits anthocyanin synthesis	Cavallini et al.,2015;Pérez-Díaz et al.,2016
	MYBC2-L1	JX050227	AFX64995	VIT_ 01s0011g04760	抑制花青素和原花青素合成Inhibits the synthesis of anthocyanin and proanthocyanin	Cavallini et al.,2015
	MYBC2-L2	GQ903730	ACX50288	VIT_ 17s0000g02660	抑制花青素合成 Inhibits anthocyanin synthesis	Zhu et al.,2018
	MYBC2-L3	KM046932	AIP98385	VIT_ 14s0006g01620	抑制花青素和原花青素合成Inhibits the synthesis of anthocyanin and proanthocyanin	Cavallini et al.,2015
	MYB86	XM_ 002280991	XP_ 002281027	VIT_ 207s0005g02480	上调LAR表达来促进原花青素合成,下调ANS和UFGT的表达来抑制花青素生物合成Activates the expression of LAR to promote proanthocyanin synthesis and suppresses the expression of ANS and UFGT to inhibit anthocyanin biosynthesis	Cheng et al.,2021
bHLH	MYC1	EU447172	ACC68685	VIT_ 07s0104g00090	与MYB5a、MYB5b、MYBA1/A2、MYBPA1互作,激活UFGT、ANR、CHI,促进花青素和原花青素合成 Interacts with MYB5a,MYB5b,MYBA1/A2,MYBPA1 and activates the expression of UFGT,ANR and CHI,to promote the synthesis of anthocyanin and procyanidin	Hichri et al.,2010
	MYCA1	ABM92332	EF193002	VIT_ 15s0046g02560	可能参与花青素和原花青素合成调控Possibility regulates the synthesis of anthocyanin and proanthocyanin	Matus et al.,2010
WD40	WDR1	ABF66625	DQ517913	VIT_ 16s0098g00870	可能参与花青素合成调控Possibility regulates the synthesis of anthocyanin	Matus et al.,2010
	WDR2	ABF66626	DQ517914	VIT_ 14s0068g00660	未知Unknown	Matus et al.,2010
WKRY	WKRY26	KF356359	AGX85877	VIT_ 08s0040g03070	与MYB5a互作,激活CHI以及液泡酸化相关基因PH1和PH5表达,参与调节液泡运输和类黄酮生物合成,在原花青素沉积中起特殊作用Interacts with MYB5a,activates the expression of CHI,PH1 and PH5 genes related to vacuole acidification,and participates in the regulation of vacuole transport and flavonoid biosynthesis,playing a special role in procyanidin deposition	Amato et al.,2017
bZIP	HY5	KF356359	AGX85877	VIT_ 04s0008g05210	响应UVB,参与黄酮醇合成调控 Responses to UVB and participates in the regulation of flavonol synthesis	Loyola et al. 2016
	HYH	KJ423106	AHX24181	VIT_ 05s0020g01090	响应UVB,参与黄酮醇合成调控 Responses to UVB and participates in the regulation of flavonol synthesis	Loyola et al. 2016
	bZIPC22	KX073969	ANA53138	VIT_ 07s0005g01450	响应UVB,参与黄酮醇合成调控 Responses to UVB and participates in the regulation of flavonol synthesis	Malacarne et al.,2016
BBX	BBX44	XM_ 002262959	XP_002262995	VIT_ 00s0347g00030	抑制HY5表达,抑制光诱导的花青苷合成Suppresses the expression of HY5 to inhibit light-induced anthocyanin synthesis	Liu et al.,2021