植物类黄酮的生物合成及其抗逆作用机制研究进展

doi:10.16420/j.issn.0513-353x.2021-1186

摘要/Abstract

摘要：

类黄酮是一类具有“黄烷”骨架的数千种衍生产物的次生代谢物质，其生物合成通路错综复杂，部分通路和相关蛋白酶已被解析和鉴定。类黄酮在植物的生长发育、花和果实的品质形成中起到重要作用。研究表明，植物类黄酮在各类胁迫下响应，有效提高了植物应对胁迫的耐性和抗性。本文综述了类黄酮在植物中的生物合成通路和分子调控机制，类黄酮在植物应对不同胁迫时的响应和作用机制，以及类黄酮未来研究的发展方向，以期为定向培育高抗性、具备深加工和产品开发的园艺作物品种提供理论支撑。

关键词: 类黄酮, 生物合成, 调控, 胁迫, 作用机制

Abstract:

Flavonoids are a class of secondary metabolites with ‘flavan’ skeleton and thousands of derivatives. The biosynthetic pathways of flavonoids are complicated，and some of them and related enzymes have been analyzed. Flavonoids play important roles in the growth and development of plants and the formation of flowers and fruits. Studies revealed that flavonoids can response to stresses and improve the tolerance and resistance of plants. This article reviews the biosynthetic pathways and molecular regulation mechanism of flavonoids in plants，the response of flavonoid pathways to different stresses，the main roles and mechanisms of flavonoids，and the future research directions. Thus，this article is expected to providing theoretical support for the targeted cultivation of horticultural crop varieties with highly resistant，deep processing，and product development.

Key words: flavonoids, biosynthesis, regulation, stress, role and mechanism

中图分类号:

Q946.8

葛诗蓓, 张学宁, 韩文炎, 李青云, 李鑫. 植物类黄酮的生物合成及其抗逆作用机制研究进展[J]. 园艺学报, 2023, 50(1): 209-224.

GE Shibei, ZHANG Xuening, HAN Wenyan, LI Qingyun, LI Xin. Research Progress on Plant Flavonoids Biosynthesis and Their Anti-stress Mechanism[J]. Acta Horticulturae Sinica, 2023, 50(1): 209-224.

图/表 5

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编号 No.	酶 Enzyme	缩写 Abbreviation	产物 Production
1	苯丙氨酸解氨酶 Phenylalanine ammonia lyase	PAL	肉桂酸 Cinnamic acid
2	肉桂酸4-羟化酶 Cinnamic acid 4-hydroxylase	C4H	p-香豆酸 p-Coumaric acid
3	4-香豆酰辅酶A连接酶 4-Coumarinyl-CoA ligase	4CL	p-香豆酰辅酶；咖啡酰辅酶A p-Coumarinyl-CoA；Caffeoyl-CoA
4	查尔酮合成酶 Chalcone synthase	CHS	柚皮素查尔酮 Naringenin chalcone
5	查尔酮异构酶 Chalcone isomerase	CHI	柚皮素 Naringenin
6	异黄酮合酶 Isoflavone synthase	IFS	异黄酮 Isoflavones
7	黄酮合酶 Flavone synthase	FS	黄酮 Flavones
8	黄烷酮3-羟化酶Flavanone 3-hydroxylase	F3H	二氢山萘酚 Dihydrokaempferol
9	黄酮3′-羟化酶 Flavonoid 3′-hydroxylase	F3'H	二氢槲皮素 Dihydroquercetin
10	黄酮3′5′-羟化酶 Flavonoid 3′5′-hydroxylase	F3'5'H	二氢杨梅素 Dihydromyricetin
11	二氢黄酮醇还原酶 Dihydroflavonol reductase	DFR	无色天竺葵素；无色矢车菊素；无色飞燕草素 Leucopelargonidin；Leucocyanidin；Leucodelphinidin
12	花青素合成酶 Anthocyanin synthase	ANS	天竺葵素；矢车菊素；飞燕草素 Pelargonidin；Cyanidin；Delphinidin
13	花青素还原酶 Afsnthocyanidin reductase	ANR	表儿茶素；表没食子酸儿茶素 (-)-Epicatechin；(-)-Epigallocatechin
14	无色花色素还原酶 Leucoanthocyanidin reductase	LAR	儿茶素；没食子酸儿茶素 (+)-Catechin；(+)-Gallocatechin
15	羟基肉桂酰基转移酶 Hydroxycinnamoyl acyltransferase	HCT	香豆酸；莽草酸 Coumarinic acid；Shikimate acid
16	p-香豆酸3羟化酶 p-Coumaric acid 3 hydroxylase	C3H	香豆酰莽草酸酯 Coumaroyl shikimate
17	咖啡酰莽草酸酯酶 Caffeoyl shikimate esterase	CSE	咖啡酸 Caffeic acid
18	咖啡酰辅酶A-O-甲基转移酶 Caffeoyl-CoA-O-methyltransferase	CCoAOMT	阿魏酰辅酶A Feruloyl-CoA
19	香豆酰辅酶A还原酶 Coumarin-CoA reductase	CCR	对香豆醛；松柏醛 p-Coumaraldehyde；Coniferyl aldehyde
20	肉桂醇脱氢酶Cinnamyl alcohol dehydrogenase	CAD	对香豆醇；松柏醇 p-Coumaryl alcohol；Coniferyl alcohol
21	阿魏酸5-羟化酶 Ferulic acid 5-hydroxylase	F5H	5-羟基松香醇 5-Hydroxyconigeryl alcohol
22	咖啡酸O-甲基转移酶 Caffeic acid O-methyltransferase	COMT	芥子醇 Sinapyl alcohol
23	黄酮醇合成酶 Flavonol synthetase	FLS	黄酮醇 Flavonols

编号 No.	酶 Enzyme	缩写 Abbreviation	产物 Production
1	苯丙氨酸解氨酶 Phenylalanine ammonia lyase	PAL	肉桂酸 Cinnamic acid
2	肉桂酸4-羟化酶 Cinnamic acid 4-hydroxylase	C4H	p-香豆酸 p-Coumaric acid
3	4-香豆酰辅酶A连接酶 4-Coumarinyl-CoA ligase	4CL	p-香豆酰辅酶；咖啡酰辅酶A p-Coumarinyl-CoA；Caffeoyl-CoA
4	查尔酮合成酶 Chalcone synthase	CHS	柚皮素查尔酮 Naringenin chalcone
5	查尔酮异构酶 Chalcone isomerase	CHI	柚皮素 Naringenin
6	异黄酮合酶 Isoflavone synthase	IFS	异黄酮 Isoflavones
7	黄酮合酶 Flavone synthase	FS	黄酮 Flavones
8	黄烷酮3-羟化酶Flavanone 3-hydroxylase	F3H	二氢山萘酚 Dihydrokaempferol
9	黄酮3′-羟化酶 Flavonoid 3′-hydroxylase	F3'H	二氢槲皮素 Dihydroquercetin
10	黄酮3′5′-羟化酶 Flavonoid 3′5′-hydroxylase	F3'5'H	二氢杨梅素 Dihydromyricetin
11	二氢黄酮醇还原酶 Dihydroflavonol reductase	DFR	无色天竺葵素；无色矢车菊素；无色飞燕草素 Leucopelargonidin；Leucocyanidin；Leucodelphinidin
12	花青素合成酶 Anthocyanin synthase	ANS	天竺葵素；矢车菊素；飞燕草素 Pelargonidin；Cyanidin；Delphinidin
13	花青素还原酶 Afsnthocyanidin reductase	ANR	表儿茶素；表没食子酸儿茶素 (-)-Epicatechin；(-)-Epigallocatechin
14	无色花色素还原酶 Leucoanthocyanidin reductase	LAR	儿茶素；没食子酸儿茶素 (+)-Catechin；(+)-Gallocatechin
15	羟基肉桂酰基转移酶 Hydroxycinnamoyl acyltransferase	HCT	香豆酸；莽草酸 Coumarinic acid；Shikimate acid
16	p-香豆酸3羟化酶 p-Coumaric acid 3 hydroxylase	C3H	香豆酰莽草酸酯 Coumaroyl shikimate
17	咖啡酰莽草酸酯酶 Caffeoyl shikimate esterase	CSE	咖啡酸 Caffeic acid
18	咖啡酰辅酶A-O-甲基转移酶 Caffeoyl-CoA-O-methyltransferase	CCoAOMT	阿魏酰辅酶A Feruloyl-CoA
19	香豆酰辅酶A还原酶 Coumarin-CoA reductase	CCR	对香豆醛；松柏醛 p-Coumaraldehyde；Coniferyl aldehyde
20	肉桂醇脱氢酶Cinnamyl alcohol dehydrogenase	CAD	对香豆醇；松柏醇 p-Coumaryl alcohol；Coniferyl alcohol
21	阿魏酸5-羟化酶 Ferulic acid 5-hydroxylase	F5H	5-羟基松香醇 5-Hydroxyconigeryl alcohol
22	咖啡酸O-甲基转移酶 Caffeic acid O-methyltransferase	COMT	芥子醇 Sinapyl alcohol
23	黄酮醇合成酶 Flavonol synthetase	FLS	黄酮醇 Flavonols

胁迫 Stress	物种 Species	类黄酮代谢途径的响应 Reseponse of flavonoid pathway	参考文献 Reference
干旱 Drought	普通小麦 Triticum aestivum	总酚、黄酮和花青素增加;CHS、CHI、F3H、FNS、FLS、DFR、ANS上调表达The contents of total phenols，flavonoids and anthocyanins，as well as the gene expression of CHS，CHI，F3H，FNS，FLS，DFR and ANS increased	Ma et al.，2014 Salehe & Barat-Ali，2020
	拟南芥 Arabidopsis	类黄酮合成通路转录因子异源表达显著增强抗旱能力 Heterologous expressing of transcription factors related to flavonoid synthesis pathway significantly enhanced drought resistance of plants	Muhammad et al.，2020
	大白菜Brassica rapa	总酚和黄酮含量增加The contents of total phenols and flavonoids increased	Rayhan & Baek，2020
	草莓 Fragaria × ananassa	PAL、C4H、4CL、DFR、ANS、FLS和UFGT基因转录水平升高 The expression of PAL，C4H，4CL，DFR，ANS，FLS and UFGT increased	Perin et al.，2019
胁迫 Stress	物种 Species	类黄酮代谢途径的响应 Reseponse of flavonoid pathway	参考文献 Reference
	黄瓜 Cucumis sativus	总酚、香草酸、4-羟基肉桂酸含量增加 Total phenols，vanillic acid and 4-hydroxycinnamic acid increased	Li et al.，2018
	百里香 Thymus vulgaris	总黄酮和多酚类物质含量增加 The contents of total flavonoids and polyphenols increased	Khalil et al.，2018
	葡萄 Vitis vinifera	儿茶素、表儿茶素、苹果酸、表儿茶素没食子酸酯、山奈酚苷、矢车菊苷、槲皮素苷含量增加;UFGT、CHS和F3H基因上调 The contents of catechin，epicatechin，malic acid，epicatechin gallate，kaempferol glycoside，cyanidin and quercetin glycoside，and the expression of UFGT， CHS and F3H increased	Castellarin et al.，2007 Griesser et al.，2015
盐Salt	藜麦 Chenopodium quinoa	总酚和类黄酮含量增加 The contents of total flavonoids and polyphenols increased	Aloisi et al.，2016
	烟草 Nicotiana langsdorffii	CHS基因过表达植株显著提高对盐的耐受性 Over-expression of CHS significantly improved salt tolerance of plants	Chen et al.，2019
	油橄榄 Olea europaea	总酚、甘露醇、槲皮素含量增加;PAL、C4H、4Cl、CHS和CHI转录水平升高 The contents of total polyphenols，mannitol and quercetinflavonoids，and the expression of PAL，C4H，4CL，CHS and CHI increased	Rossi et al.，2016
高温Heat	扁豆 Lens culinaris	没食子酸、水杨酸、绿原酸、阿魏酸和柚皮素含量增加 The contents of gallic acids，salicylic acid，chlorogenic acid，ferulic acid and naringenin increased	Swieca，2015
	烟草 Nicotiana langsdorffii	香豆酸、绿原酸、阿魏酸含量增加 The contents of coumaric acid，chlorogenic acid and ferulic acid increased	Ancillotti et al.，2015
	红豆 Vigna angularis	花青素和黄酮含量增加The contents of anthocyanins and flavones increased	Zlotek et al.，2015
	番茄 Solanum lycopersicon	黄酮醇含量增加 The flavonols content increased.	Martinez et al.，2016
	茶树 Camellia sinensis	儿茶素含量增加;C4H、F3H、DFR和ANR表达上调 The catechin content and the expression of C4H，F3H，DFR and ANR increased	Li et al.，2020
低温Cold	烟草 Nicotiana langsdorffii	木质素含量增加;PAL、HCT和CAD基因上调表达 The lignins content and the expression of PAL，HCT and CAD increased	Zhou et al.，2018
低温Cold	桃树 Prunus persica	儿茶素、槲皮素苷、山奈酚等多酚类含量增加;PAL、C4H、4Cl、CHI等酶活性升高 The contents of polyphenols such as catechin，quercetin and kaempferol，and the enzyme activity of PAL，C4H，4Cl and CHI increased	Wang et al.，2019
紫外辐射 UV-B	绿豆 Vigna radiata	总黄酮含量增加;PAL、CHI酶活增加 The total flavones content and the enzyme activity of PAL and CHI increased.	Goyal et al.，2014
	杨梅 Arbutus unedo	茶黄素、阿魏酸苷、胡桃苷含量增加 The contents of theaflavins，ferulide and juglandin increased	Nenadis et al.，2015
	莴苣 Lactuca sativa	类黄酮，酚酸类物质含量增加;PAL酶活提高 The contents of flavonoids and phenolic acids and the enzyme activity of PAL increased	Lee et al.，2014 Sytar et al.，2018
	葡萄 Vitis vinifera	黄烷醇、槲皮素、山奈酚、杨梅素、没食子酸等含量增加 The contents of flavanols，quercetin，kaempfero，myricetin and gallic acid increased	Berli et al.，2011 Alonso et al.，2016
	草莓 Fragaria × ananassa	山奈酚、矢车菊苷、天竺葵苷、槲皮素含量增加;CHS、CHI、FHT、DFR、FLS和UFGT基因上调表达 The contents of kaempferol，cyanidin，geranin and quercetin，and the expression of CHS，CHI，FHT，DFR，FLS and UFGT increased	Xu et al.，2017
砷As	茶树 Camellia sinensis	花青素含量增加;PAL、CHS酶活增加;PAL、CHS、CHI、F3H、DFR、ANS基因上调表达 The anthocyanins content，the enzyme activity of PAL and CHS，as well as the expression of PAL，CHS，CHI，F3H，DFR and ANS increased	Li et al.，2021
铜，铅，镉，锌，镍Cu，Pb，Cd， Zn，Ni	玉米 Zea mays	绿原酸、香草酸等多酚类物质和黄酮的含量增加 The contents of flavones and polyphenols such as chlorogenic acid and vanillic acid increased	Kisa et al.，2016 AbdElgawad et al.，2020
铝Al	荞麦 Fagopyrum esculentum	总酚、黄酮和花青素含量增加;PAL酶活性提高 The contents of total polyphenols，flavones and anthocyanins，and the enzyme activity of PAL increased	Smirnov et al.，2015
铜，铬，镉，铅 Cu，Cr，Cd，Pb	芥菜 Brassica juncea	花青素、类黄酮、多酚含量增加;PAL、CHS基因上调 The contents of anthocyanins，flavonoids and polyphenols，and the expression of PAL and CHS increased	Poonam et al.，2015 Kaur et al.，2017 Handa et al.，2019 Kohli et al.，2018
铜Cu	葡萄 Vitis vinifera	PAL、C4H、CHS、F3H、DFR转录水平升高，UFGT和ANR转录水平降低 The transcripts of PAL，C4H，CHS，F3H and DFR increased，while UFGT and ANR decreased	Leng et al.，2015

胁迫 Stress	物种 Species	类黄酮代谢途径的响应 Reseponse of flavonoid pathway	参考文献 Reference
干旱 Drought	普通小麦 Triticum aestivum	总酚、黄酮和花青素增加;CHS、CHI、F3H、FNS、FLS、DFR、ANS上调表达The contents of total phenols，flavonoids and anthocyanins，as well as the gene expression of CHS，CHI，F3H，FNS，FLS，DFR and ANS increased	Ma et al.，2014 Salehe & Barat-Ali，2020
	拟南芥 Arabidopsis	类黄酮合成通路转录因子异源表达显著增强抗旱能力 Heterologous expressing of transcription factors related to flavonoid synthesis pathway significantly enhanced drought resistance of plants	Muhammad et al.，2020
	大白菜Brassica rapa	总酚和黄酮含量增加The contents of total phenols and flavonoids increased	Rayhan & Baek，2020
	草莓 Fragaria × ananassa	PAL、C4H、4CL、DFR、ANS、FLS和UFGT基因转录水平升高 The expression of PAL，C4H，4CL，DFR，ANS，FLS and UFGT increased	Perin et al.，2019
胁迫 Stress	物种 Species	类黄酮代谢途径的响应 Reseponse of flavonoid pathway	参考文献 Reference
	黄瓜 Cucumis sativus	总酚、香草酸、4-羟基肉桂酸含量增加 Total phenols，vanillic acid and 4-hydroxycinnamic acid increased	Li et al.，2018
	百里香 Thymus vulgaris	总黄酮和多酚类物质含量增加 The contents of total flavonoids and polyphenols increased	Khalil et al.，2018
	葡萄 Vitis vinifera	儿茶素、表儿茶素、苹果酸、表儿茶素没食子酸酯、山奈酚苷、矢车菊苷、槲皮素苷含量增加;UFGT、CHS和F3H基因上调 The contents of catechin，epicatechin，malic acid，epicatechin gallate，kaempferol glycoside，cyanidin and quercetin glycoside，and the expression of UFGT， CHS and F3H increased	Castellarin et al.，2007 Griesser et al.，2015
盐Salt	藜麦 Chenopodium quinoa	总酚和类黄酮含量增加 The contents of total flavonoids and polyphenols increased	Aloisi et al.，2016
	烟草 Nicotiana langsdorffii	CHS基因过表达植株显著提高对盐的耐受性 Over-expression of CHS significantly improved salt tolerance of plants	Chen et al.，2019
	油橄榄 Olea europaea	总酚、甘露醇、槲皮素含量增加;PAL、C4H、4Cl、CHS和CHI转录水平升高 The contents of total polyphenols，mannitol and quercetinflavonoids，and the expression of PAL，C4H，4CL，CHS and CHI increased	Rossi et al.，2016
高温Heat	扁豆 Lens culinaris	没食子酸、水杨酸、绿原酸、阿魏酸和柚皮素含量增加 The contents of gallic acids，salicylic acid，chlorogenic acid，ferulic acid and naringenin increased	Swieca，2015
	烟草 Nicotiana langsdorffii	香豆酸、绿原酸、阿魏酸含量增加 The contents of coumaric acid，chlorogenic acid and ferulic acid increased	Ancillotti et al.，2015
	红豆 Vigna angularis	花青素和黄酮含量增加The contents of anthocyanins and flavones increased	Zlotek et al.，2015
	番茄 Solanum lycopersicon	黄酮醇含量增加 The flavonols content increased.	Martinez et al.，2016
	茶树 Camellia sinensis	儿茶素含量增加;C4H、F3H、DFR和ANR表达上调 The catechin content and the expression of C4H，F3H，DFR and ANR increased	Li et al.，2020
低温Cold	烟草 Nicotiana langsdorffii	木质素含量增加;PAL、HCT和CAD基因上调表达 The lignins content and the expression of PAL，HCT and CAD increased	Zhou et al.，2018
低温Cold	桃树 Prunus persica	儿茶素、槲皮素苷、山奈酚等多酚类含量增加;PAL、C4H、4Cl、CHI等酶活性升高 The contents of polyphenols such as catechin，quercetin and kaempferol，and the enzyme activity of PAL，C4H，4Cl and CHI increased	Wang et al.，2019
紫外辐射 UV-B	绿豆 Vigna radiata	总黄酮含量增加;PAL、CHI酶活增加 The total flavones content and the enzyme activity of PAL and CHI increased.	Goyal et al.，2014
	杨梅 Arbutus unedo	茶黄素、阿魏酸苷、胡桃苷含量增加 The contents of theaflavins，ferulide and juglandin increased	Nenadis et al.，2015
	莴苣 Lactuca sativa	类黄酮，酚酸类物质含量增加;PAL酶活提高 The contents of flavonoids and phenolic acids and the enzyme activity of PAL increased	Lee et al.，2014 Sytar et al.，2018
	葡萄 Vitis vinifera	黄烷醇、槲皮素、山奈酚、杨梅素、没食子酸等含量增加 The contents of flavanols，quercetin，kaempfero，myricetin and gallic acid increased	Berli et al.，2011 Alonso et al.，2016
	草莓 Fragaria × ananassa	山奈酚、矢车菊苷、天竺葵苷、槲皮素含量增加;CHS、CHI、FHT、DFR、FLS和UFGT基因上调表达 The contents of kaempferol，cyanidin，geranin and quercetin，and the expression of CHS，CHI，FHT，DFR，FLS and UFGT increased	Xu et al.，2017
砷As	茶树 Camellia sinensis	花青素含量增加;PAL、CHS酶活增加;PAL、CHS、CHI、F3H、DFR、ANS基因上调表达 The anthocyanins content，the enzyme activity of PAL and CHS，as well as the expression of PAL，CHS，CHI，F3H，DFR and ANS increased	Li et al.，2021
铜，铅，镉，锌，镍Cu，Pb，Cd， Zn，Ni	玉米 Zea mays	绿原酸、香草酸等多酚类物质和黄酮的含量增加 The contents of flavones and polyphenols such as chlorogenic acid and vanillic acid increased	Kisa et al.，2016 AbdElgawad et al.，2020
铝Al	荞麦 Fagopyrum esculentum	总酚、黄酮和花青素含量增加;PAL酶活性提高 The contents of total polyphenols，flavones and anthocyanins，and the enzyme activity of PAL increased	Smirnov et al.，2015
铜，铬，镉，铅 Cu，Cr，Cd，Pb	芥菜 Brassica juncea	花青素、类黄酮、多酚含量增加;PAL、CHS基因上调 The contents of anthocyanins，flavonoids and polyphenols，and the expression of PAL and CHS increased	Poonam et al.，2015 Kaur et al.，2017 Handa et al.，2019 Kohli et al.，2018
铜Cu	葡萄 Vitis vinifera	PAL、C4H、CHS、F3H、DFR转录水平升高，UFGT和ANR转录水平降低 The transcripts of PAL，C4H，CHS，F3H and DFR increased，while UFGT and ANR decreased	Leng et al.，2015

病原物分类 Classification of pathogen	病原物名称 Name of pathogen	物种 Species	类黄酮代谢途径的响应 Response of flavonoid pathway	参考文献 Reference
真菌 Fungus	异螺旋孢腔菌 Cochliobolus heterostrophus	高粱 Sorghum bicolor	花青素合成受到抑制 The synthesis of anthocyanins was inhibited	Lo & Nicholson，1998
	禾谷镰刀菌 Fusarium graminearum	小麦 Triticum aestivum	类黄酮积累 Flavonoids was accumulated.	Ravensdale et al.，2014
	茄镰孢菌 Nectria haematococca 大豆疫霉菌 Phytophthora sojae	豌豆Pisum sativum 紫花苜蓿Medicago sativa 苜蓿Medicago truncatula 鹰嘴豆Cicer arietinum 大豆Glycine max	豆科植物特异性合成异黄酮衍生物蝶形烷 ‘Butterfly alkane’，an isoflavone derivative，was specifically synthesized by legume	Liu et al.，2006 Subramanian et al.，2005
	链格孢霉 Alternaria alternata	草莓 Fragaria × ananassa	儿茶素在叶片中合成并抑制菌丝形成 Catechins were synthesized in leaves and inhibited mycelium formation	Yamamoto et al.，2000
	锈病真菌 Melampsora laricipopulina	杨树 Populus L.	黄烷-3-醇积累 Flavan-3-ols were accumulated	Ullah et al.，2019
	胶锈菌 Gymnosporangium yamadai Miyabe	苹果 Malus × domestica	儿茶素、花青素在病斑部积累 Catechins and anthocyanins was accumulated in disease spots	Lu et al.，2017
细菌 Bacteria	白叶枯萎病菌 Xanthomonas oryzae pv. Oryzae	水稻 Oryza sativa	芦丁处理激活了水杨酸信号通路 Exogenous rutin activated salicylic acid signaling pathway	Yang et al.，2016
细菌 Bacteria	丁香假单胞菌 Pst DC3000	拟南芥 Arabidopsis	花青素合成受到抑制;CHS、CHI、FLS下调表达 The synthesis of anthocyanins and the expression of CHS，CHI and FLS were inhibited	Saijo et al.，2009 Schenke et al.，2011
	炭疽菌 Colletotrichum gloeosporioides	高粱 Sorghum bicolor	木犀草素在高抗品种中积累 Luteolin was accumulated in high-resistant varieties	Du et al.，2010
	念珠菌 Candidatus liberibacter	甜橙 Citrus sinensis	黄酮苷、聚甲氧基黄酮、羟基肉桂酸酯在抗病品种中积累 Flavonoids，polymethoxyflavone and hydroxycinnamate were accumulated in resistant varieties	Hijaz et al.，2013
害虫 Insect	斜纹夜盗蛾 Spodoptera litura 棉铃虫Helicoverpa armigera	烟草 Nicotiana tabacum	黄酮醇在AtMYB12过表达植株中积累 Flavonols were accumulated in AtMYB12-overexpressed plants	Misra et al.，2010
害虫 Insect	大豆夜蛾 Anticarsia gemmatalis	大豆 Glycine max	槲皮素和山奈酚含量增加 Quercetin and kaempferol were accumulated	Gómez et al.，2018