Advances in the Regulation of Anthocyanin Biosynthesis by Hormone Signaling in Fruit Trees

doi:10.16420/j.issn.0513-353x.2022-1038

Acta Horticulturae Sinica ›› 2023, Vol. 50 ›› Issue (11): 2516-2536.doi: 10.16420/j.issn.0513-353x.2022-1038

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Advances in the Regulation of Anthocyanin Biosynthesis by Hormone Signaling in Fruit Trees

ZHENG Tianyi¹^,²^,³, HU Yujie¹^,²^,³, ZHANG Xuehao¹^,²^,³, LI Wanping¹^,²^,³, FANG Yulin¹^,²^,³, ZHANG Kekun¹^,²^,³^,^*(), CHEN Keqin¹^,²^,³^,^*()

¹ College of Enology，Northwest A & F University，Yangling，Shaanxi 712100，China
² Heyang Grape Experimental Station，Northwest A & F University，Heyang，Shaanxi 715300，China
³ The Wine Experimental Station in Eastern Foothill of Helan Mountain，Northwest A & F University，Yongning，Ningxia 750104，China

Received:2023-07-03 Revised:2023-08-14 Online:2023-11-25 Published:2023-11-28
Contact: ZHANG Kekun, CHEN Keqin

Abstract

Abstract:

This review summarizes the molecular mechanisms underlying the roles of abscisic acid，jasmonic acid，ethylene，auxin，cytokinin，brassinolide，gibberellin and strigolactone in regulating the anthocyanin accumulation in the fruit trees based on gene expression regulation of the anthocyanin biosynthesis pathway. The review will provide a theoretic basis for enriching the fundamental researches of anthocyanin regulation and biosynthesis in fruit trees，understanding the critical signal transduction pathways of plant hormones，and rationally applying plant hormones to regulate fruits development. The information provides references for the further study of the nthocyanin metabolic pathways in different types of fruit trees.

Key words: fruit trees, anthocyanin, biosynthesis, hormone

ZHENG Tianyi, HU Yujie, ZHANG Xuehao, LI Wanping, FANG Yulin, ZHANG Kekun, CHEN Keqin. Advances in the Regulation of Anthocyanin Biosynthesis by Hormone Signaling in Fruit Trees[J]. Acta Horticulturae Sinica, 2023, 50(11): 2516-2536.

Figures/Tables 3

References 148

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果树 Fruit tree	MYB	bHLH	WD40	其他 Others	参考文献 Reference
苹果Apple	MdMYB1，MdMYB3 MdMYB9，MdMYB10 MdMYB11，MdMYB12 MdMYB16，MdMYB17 MdMYB22，MdMYB60 MdMYB111，MdMYB24L	MdbHLH3，MdbHLH33 MdMYC2	MdTTG1	MdERF1B，MdERF3，MdEIL1，MdBT2，MdJa2，MdbZIP44， MdERF13，MdERF38，MdJAZ8，MdJAZ11，MdBEH2.2，MdABI5，MdTCP46，MdBBX22，MdWRKY40	An et al.,2012,2018b,2020,2021；Sun et al.,2017；Wang et al.,2019,2021,2022b；Yang et al.,2021；Li et al.,2022
梨Pear	PyMYB10，PyMYB10b PyMYB14，PyMYB114	PybHLH3	PyWD40	PyERF3，PyWRKY26 PpEIN3，PpEIL PpEIN2，PbERF22	Yao et al.,2017；Wu et al.,2020；Cui et al.,2021；Liu et al.,2021a
葡萄Grape	VvMYBA1，VvMYBA2 VvMYBA1-2，VvMYBA1-3，VvMYBA5，VvMYBA6， VvMYBA7，VvMYBCS1	VvbHLH79，VvMYC1 VvMYCA1	VvWDR1	VvSPL9，VIPYL1	Zhao et al.,2019；Kanzaki et al.,2020；Jiu et al.,2021；Su et al.,2021；Mhetre et al.,2022；Ju et al.,2022；Wang et al.,2022a；Xie et al,2022
猕猴桃Kiwifruit	AcMYB110，AcMYBC1 AcMYB75	AcbHLH1，AcbHLH4 AcbHLH5	AcWDR1	AcWRKY44	Li et al.,2017；Peng et al.,2020；Liu et al.,2021b
樱桃Cherry	PaMYB10.1-3，PavMYB10.1，PacMYBA	PabHLH3，PabHLH33	PaWD40	PacNCED1，PaMADS7 PavRaint	Qi et al.,2011；Shen et al.,2014；Starkevič et al.,2015；李丽仙等,2022
李Plum	PsMYB10.1，PsMYB73 PsMYB75，PsMYB124 PsAS1，PaKUA1	PsbHLH3，PsbHLH30 PsbHLH48，PsbHLH62 PsbHLH7，PsbHLH130 PsbHLH144，PsILR3 PsMYC2	PsCIA1，PsPUB26	PsERS1，PsETR1 PsEFR1a，PsEFR1b PsEFR2a，PsEFR3a PsEFR3b	Cheng et al.,2016；Chen et al.,2021；Fang et al.,2021
桃Peach	PpMYB10.1，PpMYB10.3 PpMYB16，PpMYB111	PpbHLH3，PpbHLH33	PpWD40	PpSPL1，PpNAC1	Gou et al.,2011；Rahim et al.,2014；Zhou et al.,2015；Sun et al.,2017
杧果Mango	MiMYB1，MinMYB10 MinMYBPA	MibHLH2		MiWRKY1，MiWRKY81	Kanzaki et al.,2020；Shi et al.,2022；Zhang et al.,2022b
草莓Strawberry	FaMYB1，FaMYB9 FaMYB10，FaMYB11 FaMYB75，FaMYB113	FabHLH3，FabHLH25 FabHLH29，FabHLH33 FabHLH80，FabHLH98	FaTTG1	FaSCL8，FaRAV1	Schaart et al.,2013；Sun et al.,2017；Karppinen et al.,2018；Zhang et al.,2020；Chen et al.,2022b
石榴 Pomegranate	PgAn2	PgAn1	PgWD40		Ben-Simhon et al.,2011
荔枝Litchi	LcMYB1，LcMYB5 LcR1MYB ，LcMYBx	LcbHLH1，LcbHLH3		LcNAC13， LcNAC90 LcSPL1	Lai et al.,2016；Liu et al.,2017；Jiang et al.,2019；Qu et al.,2022；Zhang et al.,2022b

果树 Fruit tree	MYB	bHLH	WD40	其他 Others	参考文献 Reference
苹果Apple	MdMYB1，MdMYB3 MdMYB9，MdMYB10 MdMYB11，MdMYB12 MdMYB16，MdMYB17 MdMYB22，MdMYB60 MdMYB111，MdMYB24L	MdbHLH3，MdbHLH33 MdMYC2	MdTTG1	MdERF1B，MdERF3，MdEIL1，MdBT2，MdJa2，MdbZIP44， MdERF13，MdERF38，MdJAZ8，MdJAZ11，MdBEH2.2，MdABI5，MdTCP46，MdBBX22，MdWRKY40	An et al.,2012,2018b,2020,2021；Sun et al.,2017；Wang et al.,2019,2021,2022b；Yang et al.,2021；Li et al.,2022
梨Pear	PyMYB10，PyMYB10b PyMYB14，PyMYB114	PybHLH3	PyWD40	PyERF3，PyWRKY26 PpEIN3，PpEIL PpEIN2，PbERF22	Yao et al.,2017；Wu et al.,2020；Cui et al.,2021；Liu et al.,2021a
葡萄Grape	VvMYBA1，VvMYBA2 VvMYBA1-2，VvMYBA1-3，VvMYBA5，VvMYBA6， VvMYBA7，VvMYBCS1	VvbHLH79，VvMYC1 VvMYCA1	VvWDR1	VvSPL9，VIPYL1	Zhao et al.,2019；Kanzaki et al.,2020；Jiu et al.,2021；Su et al.,2021；Mhetre et al.,2022；Ju et al.,2022；Wang et al.,2022a；Xie et al,2022
猕猴桃Kiwifruit	AcMYB110，AcMYBC1 AcMYB75	AcbHLH1，AcbHLH4 AcbHLH5	AcWDR1	AcWRKY44	Li et al.,2017；Peng et al.,2020；Liu et al.,2021b
樱桃Cherry	PaMYB10.1-3，PavMYB10.1，PacMYBA	PabHLH3，PabHLH33	PaWD40	PacNCED1，PaMADS7 PavRaint	Qi et al.,2011；Shen et al.,2014；Starkevič et al.,2015；李丽仙等,2022
李Plum	PsMYB10.1，PsMYB73 PsMYB75，PsMYB124 PsAS1，PaKUA1	PsbHLH3，PsbHLH30 PsbHLH48，PsbHLH62 PsbHLH7，PsbHLH130 PsbHLH144，PsILR3 PsMYC2	PsCIA1，PsPUB26	PsERS1，PsETR1 PsEFR1a，PsEFR1b PsEFR2a，PsEFR3a PsEFR3b	Cheng et al.,2016；Chen et al.,2021；Fang et al.,2021
桃Peach	PpMYB10.1，PpMYB10.3 PpMYB16，PpMYB111	PpbHLH3，PpbHLH33	PpWD40	PpSPL1，PpNAC1	Gou et al.,2011；Rahim et al.,2014；Zhou et al.,2015；Sun et al.,2017
杧果Mango	MiMYB1，MinMYB10 MinMYBPA	MibHLH2		MiWRKY1，MiWRKY81	Kanzaki et al.,2020；Shi et al.,2022；Zhang et al.,2022b
草莓Strawberry	FaMYB1，FaMYB9 FaMYB10，FaMYB11 FaMYB75，FaMYB113	FabHLH3，FabHLH25 FabHLH29，FabHLH33 FabHLH80，FabHLH98	FaTTG1	FaSCL8，FaRAV1	Schaart et al.,2013；Sun et al.,2017；Karppinen et al.,2018；Zhang et al.,2020；Chen et al.,2022b
石榴 Pomegranate	PgAn2	PgAn1	PgWD40		Ben-Simhon et al.,2011
荔枝Litchi	LcMYB1，LcMYB5 LcR1MYB ，LcMYBx	LcbHLH1，LcbHLH3		LcNAC13， LcNAC90 LcSPL1	Lai et al.,2016；Liu et al.,2017；Jiang et al.,2019；Qu et al.,2022；Zhang et al.,2022b

激素 Hormone	果树品种 Cultivar	处理时期 Stage	处理 Treatment	总花色苷含量 Anthocyanin content	相关基因 Related gene	参考文献 Reference
脱落酸 Abscisic acid	葡萄：火焰无核 Grape： Flame Seedless	转色期 Veraison	400 mg · L^-1 ABA喷施植株 Spraying plants with 400 mg · L^-1 of ABA	增加 Increase	F3’H ↑ F3’5’H ↑ DFR ↑ LDOX ↑ UFG ↑ TOMT ↑	Mhetre et al.,2022
	草莓：章姬 Strawberry： Akihime	大绿期 Large green stage	1.0 mmol · L^-1 ABA浸泡草莓果实5min Imersing strawberrie fruit in 1.0 mmol · L^-1 abscisic acid for 5 min	增加 Increase		Li et al.,2014
	荔枝：桂味 Litchi：Guiwei	商业成熟期 Commercial maturity	0.1 g · L^-1ABA浸泡荔枝果实1 min Imersing litch fruit in 0.1 g · L^-1ABA for 1 min	增加 Increase	PAL ↑ C4H ↑ 4CL ↑ CHS ↑ F3H ↑ ANS ↑ UFGT ↑	Qu et al.,2022
	欧洲越橘 Bilberry	未成熟时期 Unripe green stage	0.5或2 mmol · L^-1ABA混合0.5% Tween20喷施未成熟的绿色越橘果实 Spraying immature green bilberry fruit with 0.5 or 2 mmol · L^-1 ABA solution mixed with 0.5% Tween20	增加 Increase	CHS ↑ ANS ↑ UFGT ↑
茉莉酸 Jasmonic acid	桃：白凤 Peach：Baifeng	商业成熟期（盛花期后100 d） Commercial maturity（approximately 100 d after full bloom）	含200 μmol · L^-1茉莉酸的0.1%乙醇溶液中浸泡果实15 min Imersing peach fruit in 200 μmol · L^-1 jasmonic acid mixed with 0.1% ethanol for 15 min	增加Increase	PAL ↑ CHS ↑ CHI ↑ F3H ↑ DFR ↑ ANS ↑ UFGT ↑ MYB10.1 ↑ bHLH3 ↑ WD40 ↑	Tang et al.,2022
	红肉苹果 Red-fleshed apple		10^-4 mol · L^-1茉莉酸甲酯处理红肉苹果愈伤组织 Culturing apple callus on medium containing 10^-4 mol · L^-1 methyl jasmonate	增加Increase	CHI ↑ CHS ↑ DFR ↑ F3H ↑ LDOX ↑ UFGT ↑ MYB3 ↑ MYB9 ↑ MYB10 ↑ MYB17 ↑ MYB111 ↑ MYB11 ↓	Sun et al.,2017
	覆盆子：秋季极乐 Redraspberry： Autumn Bliss	白果期 White stage	100 μmol · L^-1 茉莉酸甲酯喷施果实 Spraying fruit with 100 μmol · L^-1 methyl jasmonate	增加Increase	ANS ↑ DFR ↑ MYB10 ↑	Moro et al.,2017
乙烯Ehylene	葡萄：火焰无核 Grape： Flame Seedless	转色期 Veraison	400 mg · L^-1乙烯利喷施植株 Spraying plants with 400 mg · L^-1of ethephon	增加Increase	F3’H ↑ F3’5’H ↑ DFR ↑ LDOX ↑ UFGT ↑ OMT ↑	Mhetre et al.,2022
	葡萄：藤稔 Grape： Fujiminori	转色前 Pre-veraison stage	500 mg · L^-1乙烯利喷施植株 Spraying grapes with 500 mg · L^-1 of ethephon	增加Increase	PAL ↑ C4H ↑ CHS ↑ CHI ↑ F3H ↑ UFGT ↑ MYBA1 ↑ UFGT ↑ LDOX ↑ MYBA2 ↑ MYBA3 ↑	Wang et al.,2022a
	苹果：早捷 Apple： Geneva Early	成熟期（开花后60 d） Maturity (60 d after full bloom)	1 000 mg · L^-1乙烯利浸泡苹果果实1 min Imersing apple fruit in 1 000 mg · L^-1 ethephon solution for 1 min	增加Increase	MYC2 ↑	Wang et al.,2022b
	杧果：贵妃 Mango：Guifei	绿熟期 Mature green stage	900 mg · L^-1乙烯利浸泡杧果果实10 min Imersing mango fruit in 900 mg · L^-1 ethephon solution for 10 min	增加Increase		Chen et al.,2022a
生长素 Auxin	红肉苹果 Red-fleshed apple		在分别含有5、10、20、40 μmol · L^-1NAA的培养基上培养苹果愈伤组织 Culturing apple callus on medium containing 5, 10, 20 or 40 μmol · L^-1NAA respectively	下降Decrease	CHS ↓ CHI ↓ DFR ↓ MYB3 ↓ MYB9 ↓ MYB10 ↓ bHLH3 ↓ bHLH33 ↓	Wang et al.,2018b
	桃：大久保 Peach：Dajiubao	开花后70 d 70 d after flowering	0.1或1 mmol · L^-1IAA溶液浸泡果实10 min Imersing fruit in 0.1 or 1 mmol · L^-1 IAA solution for 10 min	增加Increase	CHS ↑ DFR ↑ F3H ↑ UFGT↑	Liu,2019
	葡萄：温克 Grape：Wink	转色前1周One week before color conversion	200 mg · L^-1 NAA喷施葡萄果实 Spraying grape berries with 200 mg · L^-1 of NAA		C4L ↓ CHS ↓ CHI ↓ F3H ↓ LDOX ↓ UFGT ↓ OMT ↓	Su et al.,2021
	覆盆子：秋季极乐 Red raspberry： Autumn Bliss	白果期 White stage	100 μmol · L^-1 IAA溶液喷施果实 Spraying grape berries with 100 μmol · L^-1 of IAA	下降Decrease	ANS ↓ MYB10 ↓	Moro et al.,2017
油菜素内酯 Brassinolide	葡萄：克瑞森无核Grape：Crimson Seedless	转色期 Veraison	1.0 或2.0 mg · L^-1油菜素内酯喷施植株 Spraying plants with 1.0 or 2.0 mg · L^-1brassinolide	增加Increase		Belal et al.,2022
	葡萄：赤霞珠 Grape： Cabernet Sauvignon：	转色前 Pre-veraison stage	0.1、0.4和0.8 mg · L^-1EBR喷施植株 Spraying pants with 0.1，0.4 or 0.8 mg · L^-1 EBR	增加Increase	CHS ↑ F3H ↑ F3’H ↑ F3’5’H ↑ DFR ↑ CHI ↓ ANS ↓ UFGT ↓	栾丽英,2014
	苹果：红田 Apple：Red field		1 μmol · L^-1 BL处理培养基中苹果幼苗 Culturing apple seedlings on medium containing 1 μmol · L^-1 BL	下降Decrease	F3H ↓ ANS ↓ UFGT ↓ CHS ↓ DFR ↓ MYB9 ↓ MYB11 ↓ MYB22 ↓ bHLH3 ↓ bHLH33↓	Wang et al.,2021
细胞分裂素 Cytokinin	红肉苹果 Red-fleshed apple		含4 mmol · L^-1 6-BA的培养基分别培养苹果愈伤组织3、6、12、24 h Culturing apple callus on medium containing 4 mmol · L^-16-BA for 3，6，12 or 24 hours，respectively	增加Increase	CHS ↑ CHI ↑ F3H ↑ DFR ↑ LDOX ↑ UFGT ↑	Wang et al.,2018a
	草莓：章姬Strawberry： Akihime	开花后7 d One week after anthesis	5、10、15 mg · L^-1CPPU喷施植株 Spraying plants with 5，10 or 15 mg · L^-1 CPPU	增加Increase		Li et al.,2016
	荔枝：妃子笑 Litchi： Feizixiao	变色期 Color break stage	4 mg · L^-1CPPU喷施果簇 Spraying fruit clusters with 4 mg · L^-1CPPU	下降Decrease	PAL ↓ C4H ↓ CHS ↓ LDOX ↓	Hu et al.,2018
	草莓：Ruegun Strawberry： Ruegun	人工授粉后1，3，6，7，9 d 1,3,6,7,9 d after pollination	200 μmol · L^-1 6-BA喷施植株 Spraying plants with 200 μmol · L^-1 6-BA	增加Increase		原乔慧,2019
赤霉素 Gibberellin	葡萄：Skookum 无核Grape：Skookum Seedless	开花前12 d，开花后12 d，开花后21 d 12 d before flowering;12, 21 d after flowering	15和30 mg · L^-1 GA₃喷施植株 Spraying plants with 15 or 30 mg · L^-1 GA₃	下降Decrease		Reynolds et al.,2016
	葡萄：西拉 Grape： Syrah	盛花前20和15 d 20 d and 15 d before full bloom	将花序分别浸入含有3、5、7 mg · L^-1GA₃的0.1% Tween溶液中 Dipping inflorescence bunch in 0.1% Tween solution containing 3，5，or 7 mg · L^-1GA₃ respectively	增加Increase		Xie et al.,2022
	蓝莓：安娜 Blueberry： Anna	完全坐果后 After the plants fully set fruit	15和30 mg · L^-1 GA₃喷施植株 Spraying plants with 15 or 30 mg · L^-1 GA₃	增加Increase		杨海燕等,2020
	荔枝：桂味 Litchi： Guiwei	商业成熟期Commercial maturity	0.05 g · L^-1GA₃浸泡荔枝果实1 min Imersing litch fruit in 0.05 g · L^-1GA₃ for 1 min	增加Increase	PAL ↑ C4H ↑ 4CL ↑ CHS ↑ ANS ↑	Qu et al.,2022
独角金内酯 Strigolactone	葡萄：赤霞珠 Grape： Cabernet Sauvignon	转色前期 At the start of veraison	10 μmol · L^-1 独脚金内酯类似物 rac-GR24喷施植株，7 d后再次喷施 Spraying plants with 10 μmol · L^-1 rac-GR24（The SL analogue）. Seven days later，second treatments were made using the same concentration and method	增加Increase	PAL ↑ F3’5’H ↑ LDOX ↑ DFR ↑	Ju et al.,2022
	葡萄：巴贝拉 Grape： Barbera	转色期 Veraison	10^-5 mol · L^-1 独脚金内酯类似物rac-GR24喷施植株 Spraying plants with 10^-5 mol · L^-1 rac-GR24（The SL analogue）	略有下降Decrease lightly	UFGT— MYB1—	Ferrero et al.,2018

激素 Hormone	果树品种 Cultivar	处理时期 Stage	处理 Treatment	总花色苷含量 Anthocyanin content	相关基因 Related gene	参考文献 Reference
脱落酸 Abscisic acid	葡萄：火焰无核 Grape： Flame Seedless	转色期 Veraison	400 mg · L^-1 ABA喷施植株 Spraying plants with 400 mg · L^-1 of ABA	增加 Increase	F3’H ↑ F3’5’H ↑ DFR ↑ LDOX ↑ UFG ↑ TOMT ↑	Mhetre et al.,2022
	草莓：章姬 Strawberry： Akihime	大绿期 Large green stage	1.0 mmol · L^-1 ABA浸泡草莓果实5min Imersing strawberrie fruit in 1.0 mmol · L^-1 abscisic acid for 5 min	增加 Increase		Li et al.,2014
	荔枝：桂味 Litchi：Guiwei	商业成熟期 Commercial maturity	0.1 g · L^-1ABA浸泡荔枝果实1 min Imersing litch fruit in 0.1 g · L^-1ABA for 1 min	增加 Increase	PAL ↑ C4H ↑ 4CL ↑ CHS ↑ F3H ↑ ANS ↑ UFGT ↑	Qu et al.,2022
	欧洲越橘 Bilberry	未成熟时期 Unripe green stage	0.5或2 mmol · L^-1ABA混合0.5% Tween20喷施未成熟的绿色越橘果实 Spraying immature green bilberry fruit with 0.5 or 2 mmol · L^-1 ABA solution mixed with 0.5% Tween20	增加 Increase	CHS ↑ ANS ↑ UFGT ↑
茉莉酸 Jasmonic acid	桃：白凤 Peach：Baifeng	商业成熟期（盛花期后100 d） Commercial maturity（approximately 100 d after full bloom）	含200 μmol · L^-1茉莉酸的0.1%乙醇溶液中浸泡果实15 min Imersing peach fruit in 200 μmol · L^-1 jasmonic acid mixed with 0.1% ethanol for 15 min	增加Increase	PAL ↑ CHS ↑ CHI ↑ F3H ↑ DFR ↑ ANS ↑ UFGT ↑ MYB10.1 ↑ bHLH3 ↑ WD40 ↑	Tang et al.,2022
	红肉苹果 Red-fleshed apple		10^-4 mol · L^-1茉莉酸甲酯处理红肉苹果愈伤组织 Culturing apple callus on medium containing 10^-4 mol · L^-1 methyl jasmonate	增加Increase	CHI ↑ CHS ↑ DFR ↑ F3H ↑ LDOX ↑ UFGT ↑ MYB3 ↑ MYB9 ↑ MYB10 ↑ MYB17 ↑ MYB111 ↑ MYB11 ↓	Sun et al.,2017
	覆盆子：秋季极乐 Redraspberry： Autumn Bliss	白果期 White stage	100 μmol · L^-1 茉莉酸甲酯喷施果实 Spraying fruit with 100 μmol · L^-1 methyl jasmonate	增加Increase	ANS ↑ DFR ↑ MYB10 ↑	Moro et al.,2017
乙烯Ehylene	葡萄：火焰无核 Grape： Flame Seedless	转色期 Veraison	400 mg · L^-1乙烯利喷施植株 Spraying plants with 400 mg · L^-1of ethephon	增加Increase	F3’H ↑ F3’5’H ↑ DFR ↑ LDOX ↑ UFGT ↑ OMT ↑	Mhetre et al.,2022
	葡萄：藤稔 Grape： Fujiminori	转色前 Pre-veraison stage	500 mg · L^-1乙烯利喷施植株 Spraying grapes with 500 mg · L^-1 of ethephon	增加Increase	PAL ↑ C4H ↑ CHS ↑ CHI ↑ F3H ↑ UFGT ↑ MYBA1 ↑ UFGT ↑ LDOX ↑ MYBA2 ↑ MYBA3 ↑	Wang et al.,2022a
	苹果：早捷 Apple： Geneva Early	成熟期（开花后60 d） Maturity (60 d after full bloom)	1 000 mg · L^-1乙烯利浸泡苹果果实1 min Imersing apple fruit in 1 000 mg · L^-1 ethephon solution for 1 min	增加Increase	MYC2 ↑	Wang et al.,2022b
	杧果：贵妃 Mango：Guifei	绿熟期 Mature green stage	900 mg · L^-1乙烯利浸泡杧果果实10 min Imersing mango fruit in 900 mg · L^-1 ethephon solution for 10 min	增加Increase		Chen et al.,2022a
生长素 Auxin	红肉苹果 Red-fleshed apple		在分别含有5、10、20、40 μmol · L^-1NAA的培养基上培养苹果愈伤组织 Culturing apple callus on medium containing 5, 10, 20 or 40 μmol · L^-1NAA respectively	下降Decrease	CHS ↓ CHI ↓ DFR ↓ MYB3 ↓ MYB9 ↓ MYB10 ↓ bHLH3 ↓ bHLH33 ↓	Wang et al.,2018b
	桃：大久保 Peach：Dajiubao	开花后70 d 70 d after flowering	0.1或1 mmol · L^-1IAA溶液浸泡果实10 min Imersing fruit in 0.1 or 1 mmol · L^-1 IAA solution for 10 min	增加Increase	CHS ↑ DFR ↑ F3H ↑ UFGT↑	Liu,2019
	葡萄：温克 Grape：Wink	转色前1周One week before color conversion	200 mg · L^-1 NAA喷施葡萄果实 Spraying grape berries with 200 mg · L^-1 of NAA		C4L ↓ CHS ↓ CHI ↓ F3H ↓ LDOX ↓ UFGT ↓ OMT ↓	Su et al.,2021
	覆盆子：秋季极乐 Red raspberry： Autumn Bliss	白果期 White stage	100 μmol · L^-1 IAA溶液喷施果实 Spraying grape berries with 100 μmol · L^-1 of IAA	下降Decrease	ANS ↓ MYB10 ↓	Moro et al.,2017
油菜素内酯 Brassinolide	葡萄：克瑞森无核Grape：Crimson Seedless	转色期 Veraison	1.0 或2.0 mg · L^-1油菜素内酯喷施植株 Spraying plants with 1.0 or 2.0 mg · L^-1brassinolide	增加Increase		Belal et al.,2022
	葡萄：赤霞珠 Grape： Cabernet Sauvignon：	转色前 Pre-veraison stage	0.1、0.4和0.8 mg · L^-1EBR喷施植株 Spraying pants with 0.1，0.4 or 0.8 mg · L^-1 EBR	增加Increase	CHS ↑ F3H ↑ F3’H ↑ F3’5’H ↑ DFR ↑ CHI ↓ ANS ↓ UFGT ↓	栾丽英,2014
	苹果：红田 Apple：Red field		1 μmol · L^-1 BL处理培养基中苹果幼苗 Culturing apple seedlings on medium containing 1 μmol · L^-1 BL	下降Decrease	F3H ↓ ANS ↓ UFGT ↓ CHS ↓ DFR ↓ MYB9 ↓ MYB11 ↓ MYB22 ↓ bHLH3 ↓ bHLH33↓	Wang et al.,2021
细胞分裂素 Cytokinin	红肉苹果 Red-fleshed apple		含4 mmol · L^-1 6-BA的培养基分别培养苹果愈伤组织3、6、12、24 h Culturing apple callus on medium containing 4 mmol · L^-16-BA for 3，6，12 or 24 hours，respectively	增加Increase	CHS ↑ CHI ↑ F3H ↑ DFR ↑ LDOX ↑ UFGT ↑	Wang et al.,2018a
	草莓：章姬Strawberry： Akihime	开花后7 d One week after anthesis	5、10、15 mg · L^-1CPPU喷施植株 Spraying plants with 5，10 or 15 mg · L^-1 CPPU	增加Increase		Li et al.,2016
	荔枝：妃子笑 Litchi： Feizixiao	变色期 Color break stage	4 mg · L^-1CPPU喷施果簇 Spraying fruit clusters with 4 mg · L^-1CPPU	下降Decrease	PAL ↓ C4H ↓ CHS ↓ LDOX ↓	Hu et al.,2018
	草莓：Ruegun Strawberry： Ruegun	人工授粉后1，3，6，7，9 d 1,3,6,7,9 d after pollination	200 μmol · L^-1 6-BA喷施植株 Spraying plants with 200 μmol · L^-1 6-BA	增加Increase		原乔慧,2019
赤霉素 Gibberellin	葡萄：Skookum 无核Grape：Skookum Seedless	开花前12 d，开花后12 d，开花后21 d 12 d before flowering;12, 21 d after flowering	15和30 mg · L^-1 GA₃喷施植株 Spraying plants with 15 or 30 mg · L^-1 GA₃	下降Decrease		Reynolds et al.,2016
	葡萄：西拉 Grape： Syrah	盛花前20和15 d 20 d and 15 d before full bloom	将花序分别浸入含有3、5、7 mg · L^-1GA₃的0.1% Tween溶液中 Dipping inflorescence bunch in 0.1% Tween solution containing 3，5，or 7 mg · L^-1GA₃ respectively	增加Increase		Xie et al.,2022
	蓝莓：安娜 Blueberry： Anna	完全坐果后 After the plants fully set fruit	15和30 mg · L^-1 GA₃喷施植株 Spraying plants with 15 or 30 mg · L^-1 GA₃	增加Increase		杨海燕等,2020
	荔枝：桂味 Litchi： Guiwei	商业成熟期Commercial maturity	0.05 g · L^-1GA₃浸泡荔枝果实1 min Imersing litch fruit in 0.05 g · L^-1GA₃ for 1 min	增加Increase	PAL ↑ C4H ↑ 4CL ↑ CHS ↑ ANS ↑	Qu et al.,2022
独角金内酯 Strigolactone	葡萄：赤霞珠 Grape： Cabernet Sauvignon	转色前期 At the start of veraison	10 μmol · L^-1 独脚金内酯类似物 rac-GR24喷施植株，7 d后再次喷施 Spraying plants with 10 μmol · L^-1 rac-GR24（The SL analogue）. Seven days later，second treatments were made using the same concentration and method	增加Increase	PAL ↑ F3’5’H ↑ LDOX ↑ DFR ↑	Ju et al.,2022
	葡萄：巴贝拉 Grape： Barbera	转色期 Veraison	10^-5 mol · L^-1 独脚金内酯类似物rac-GR24喷施植株 Spraying plants with 10^-5 mol · L^-1 rac-GR24（The SL analogue）	略有下降Decrease lightly	UFGT— MYB1—	Ferrero et al.,2018

Advances in the Regulation of Anthocyanin Biosynthesis by Hormone Signaling in Fruit Trees

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