激素信号调控果树果实花色苷合成机制研究进展

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

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

激素信号调控果树果实花色苷合成机制研究进展

郑天一¹^,²^,³, 胡玉杰¹^,²^,³, 张学昊¹^,²^,³, 李婉平¹^,²^,³, 房玉林¹^,²^,³, 张克坤¹^,²^,³^,^*(), 陈可钦¹^,²^,³^,^*()

¹ 西北农林科技大学葡萄酒学院，陕西杨凌 712100
² 西北农林科技大学合阳葡萄试验示范站，陕西合阳 715300
³ 西北农林科技大学宁夏贺兰山东麓葡萄酒试验示范站，宁夏永宁 750104

收稿日期:2023-07-03 修回日期:2023-08-14 出版日期:2023-11-25 发布日期:2023-11-28
通讯作者:
*（E-mail：zhangkekun1990@nwafu.edu.cn，
chenkeqin1985@nwafu.edu.cn）
基金资助:
国家自然科学基金项目(32002023); 现代农业产业技术体系建设专项(CARS-29-zp-6)

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 Published:2023-11-25 Online:2023-11-28
Contact:
*（E-mail：zhangkekun1990@nwafu.edu.cn，
chenkeqin1985@nwafu.edu.cn）

摘要/Abstract

摘要：

围绕花色苷合成路径相关基因的表达调控，综述了脱落酸、茉莉酸、乙烯、生长素、细胞分裂素、油菜素内酯、赤霉素、独脚金内酯对果树中花色苷积累的调控作用及其内在分子机制，以期为明确植物激素信号转导关键通路，合理应用植物激素类物质调控果实发育提供理论支撑，同时也为不同果树花色苷代谢通路的深入研究提供参考。

关键词: 果树, 花色苷, 生物合成, 激素

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

郑天一, 胡玉杰, 张学昊, 李婉平, 房玉林, 张克坤, 陈可钦. 激素信号调控果树果实花色苷合成机制研究进展[J]. 园艺学报, 2023, 50(11): 2516-2536.

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.

图/表 3

参考文献 148

[1]	An J P, Wang X F, Hao Y J. 2020. BTB/TAZ protein MdBT 2 integrates multiple hormonal and environmental signals to regulate anthocyanin biosynthesis in apple. Journal of Integrative Plant Biology, 62 (11):1643-1646. doi: 10.1111/jipb.v62.11 URL
[2]	An J P, Wang X F, Li Y Y, Song L Q, Zhao L L, You C X, Hao Y J. 2018a. EIN3-LIKE1,MYB1,and ETHYLENE RESPONSE FACTOR3 act in a regulatory loop that synergistically modulates ethylene biosynthesis and anthocyanin accumulation. Plant Physiology, 178 (2):808-823. doi: 10.1104/pp.18.00068 URL
[3]	An J P, Yao J F, Xu R R, You C X, Wang X F, Hao Y J. 2018b. Apple bZIP transcription factor MdbZIP 44 regulates abscisic acid‐promoted anthocyanin accumulation. Plant,Cell & Environment, 41 (11):2678-2692.
[4]	An J P, Zhang X W, Liu Y J, Wang X F, You C X, Hao Y J. 2021. ABI5 regulates ABA-induced anthocyanin biosynthesis by modulating the MYB1-bHLH3 complex in apple. Journal of Experimental Botany, 72 (4):1460-1472. doi: 10.1093/jxb/eraa525 URL
[5]	An Jian-ping. 2020. Study on the mechanism of hormonal and environmental signals regulating anthocyanin biosynthesis in apple[Ph. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	安建平. 2020. 激素和环境信号调控苹果花青苷生物合成的机理研究[博士论文]. 泰安: 山东农业大学.
[6]	An X H, Tian Y, Chen K Q, Wang X F, Hao Y J. 2012. The apple WD 40 protein MdTTG1 interacts with bHLH but not MYB proteins to regulate anthocyanin accumulation. Journal of Plant Physiology, 169 (7):710-717. doi: 10.1016/j.jplph.2012.01.015 URL
[7]	Belal B E A, El Kenawy M A, Omar A S M. 2022. Using brassinolide and girdling combined application as an alternative to ethephon for improving color and quality of‘Crimson Seedless’grapevines. Horticulture,Environment,and Biotechnology, doi:10.1007/s13580-022-00445-3.
[8]	Ben-Simhon Z, Judeinstein S, Fau-Nadler-Hassar T, Nadler-Hassar T, Fau-Trainin T, Trainin T, Fau-Bar-Yáakov I, Bar-Yáakov I, Fau- Borochov-Neori H, Borochov-Neori H Fau-Holland D, Holland D. 2011. A pomegranate(Punica granatum L.). WD40-repeat gene is a functional homologue of Arabidopsis TTG1 and is involved in the regulation of anthocyanin biosynthesis during pomegranate fruit development. Planta, 234 (5):865-881. doi: 10.1007/s00425-011-1438-4 pmid: 21643990
[9]	Bhadoria P, Nagar M, Bharihoke V, Bhadoria A S. 2018. Ethephon,an organophosphorous,a fruit and vegetable ripener:has potential hepatotoxic effects? Journal of Family Medicine and Primary Care, 7 (1):179-183. doi: 10.4103/jfmpc.jfmpc_422_16 pmid: 29915756
[10]	Bi Siqi, An Jianping, Wang Xiaofei, Hao Yujin, Rui Lin, Li Tong, Han Yuepeng, You Chunxiang. 2019. Ethylene response factor MdERF 3 promotes anthocyanin and proanthocyanidin accumulation in apple. Acta Horticulturae Sinica, 46 (12):2277-2285. (in Chinese)
	毕思琦, 安建平, 王小非, 郝玉金, 芮麟, 李彤, 韩月彭, 由春香. 2019. 苹果乙烯响应因子MdERF3促进花青苷和原花青苷积累. 园艺学报, 46 (12):2277-2285. doi: 10.16420/j.issn.0513-353x.2019-0122 URL
[11]	Bieza K, Lois R. 2001. An Arabidopsis mutant tolerant to lethal ultraviolet-B levels shows constitutively elevated accumulation of flavonoids and other phenolics. Plant Physiology, 126 (3):1105-1115. doi: 10.1104/pp.126.3.1105 pmid: 11457961
[12]	Binder B M. 2020. Ethylene signaling in plants. Journal of Biological Chemistry, 295 (22):7710-7725. doi: 10.1074/jbc.REV120.010854 pmid: 32332098
[13]	Bradow J M, Connick W J. 1988. Seed-germination inhibition by volatile alcohols and other compounds associated with Amaranthus palmeri residues. Journal of Chemical Ecology, 14 (7):1633-1648. doi: 10.1007/BF01012528 pmid: 24276435
[14]	Chen Han-lei, Bi Shu-wen, Ni Ruo-yi, Lei You-zhen, Quan Jin-e. 2022. Effects of different auxin and its mass concentration on mulberry cutting rooting. Journal of Henan Agricultural University, 56 (6):958-967.
	陈寒蕾, 毕舒雯, 倪若荑, 雷友贞, 权金娥. 2022. 不同生长素及质量浓度对桑树扦插生根的影响. 河南农业大学学报, 56 (6):958-967.
[15]	Chen L, Wang X S, Cui L, Zhang Y B. 2021. Transcriptome and metabolome analyses reveal pathways associated with fruit color in plum(Prunus salicina Lindl.). bioRxiv,doi:10.7717/peerj.14413.
[16]	Chen M, Gu H, Wang L, Shao Y, Li R, Li W. 2022a. Exogenous ethylene promotes peel color transformation by regulating the degradation of chlorophyll and synthesis of anthocyanin in postharvest mango fruit. Frontiers in Nutrition, 9:911542. doi: 10.3389/fnut.2022.911542 URL
[17]	Chen X, Song Z, Pang Q, Lv J, Yu L, Li T, Fang J, Jia H. 2022b. Effect of postharvest application of jasmonic acid on shelf-life quality of strawberries. European Journal of Horticultural Science,doi:10.17660/eJHS.2022/016.
[18]	Chen X M, Shi X Y, Ai Q, Han J Y, Wang H S, Fu Q S. 2022c. Transcriptomic and metabolomic analyses reveal that exogenous strigolactones alleviate the response of melon root to cadmium stress. Horticultural Plant Journal, 8 (5):637-649. doi: 10.1016/j.hpj.2022.07.001 URL
[19]	Cheng Y, Liu L, Yuan C, Guan J. 2016. Molecular characterization of ethylene-regulated anthocyanin biosynthesis in plums during fruit ripening. Plant Molecular Biology Reporter, 34 (4):777-785. doi: 10.1007/s11105-015-0963-x URL
[20]	Chervin C, Terrier N, Ageorges A, Ribes F, Kuapunyakoon T. 2006. Influence of ethylene on sucrose accumulation in grape berry. American Journal of Enology and Viticulture, 57 (4):511-513. doi: 10.5344/ajev.2006.57.4.511 URL
[21]	Clayton-Cuch D, Yu L, Shirley N, Bradley D, Bulone V, Böttcher C. 2021. Auxin treatment enhances anthocyanin production in the non-climacteric sweet cherry (Prunus avium L.). International Journal of Molecular Sciences, 22 (19):10760. doi: 10.3390/ijms221910760 URL
[22]	Cui D, Zhao S, Xu H, Allan A C, Zhang X, Fan L, Chen L, Su J, Shu Q, Li K. 2021. The interaction of MYB,bHLH and WD 40 transcription factors in red pear(Pyrus pyrifolia) peel. Plant Molecular Biology, 106 (4):407-417. doi: 10.1007/s11103-021-01160-w
[23]	Dai Hongjun, Wei Qiang, He Yan, Wang Yuening, Wang Zhenping. 2023. Effects of brassinolide treatment on anthocyanin synthesis and quality in grape under high temperature stress. Acta Horticulturae Sinica, 50 (8):1711-1722. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-0599 URL
	代红军, 魏强, 贺琰, 汪月宁, 王振平. 2023. 油菜素内酯对高温胁迫下葡萄花色苷合成及果实品质的影响. 园艺学报, 50 (8):1711-1722.
[24]	Davière J M, Achard P. 2013. Gibberellin signaling in plants. Development, 140 (6):1147-1151. doi: 10.1242/dev.087650 URL
[25]	Dejonghe W, Okamoto M, Cutler S R. 2018. Small molecule probes of ABA biosynthesis and signaling. Plant and Cell Physiology, 59 (8):1490-1499. doi: 10.1093/pcp/pcy126 pmid: 29986078
[26]	Dong T, Zheng T, Fu W, Guan L, Jia H, Fang J. 2020. The effect of ethylene on the color change and resistance to Botrytis cinerea infection in‘Kyoho grape’fruits. Foods, 9 (7):892. doi: 10.3390/foods9070892 URL
[27]	Dong Tian-yu. 2020. Effects of ethephon on ripening and disease resistance of grape fruit[M. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	董天宇. 2020. 乙烯利对葡萄果实成熟、抗病的效果探究[硕士论文]. 南京: 南京农业大学.
[28]	Fang Z, Lin-Wang K, Jiang C, Zhou D, Lin Y, Pan S, Espley R V, Ye X. 2021. Postharvest temperature and light treatments induce anthocyanin accumulation in peel of‘Akihime’plum (Prunus salicina Lindl.) via transcription factor PsMYB10.1. Postharvest Biology and Technology, 179:111592. doi: 10.1016/j.postharvbio.2021.111592 URL
[29]	Ferrero M, Pagliarani C, Novák O, Ferrandino A, Cardinale F, Visentin I, Schubert A. 2018. Exogenous strigolactone interacts with abscisic acid-mediated accumulation of anthocyanins in grapevine berries. Journal of Experimental Botany, 69 (9):2391-2401. doi: 10.1093/jxb/ery033 pmid: 29401281
[30]	Fujita Y, Fujita M, Shinozaki K, Yamaguchi-Shinozaki K. 2011. ABA-mediated transcriptional regulation in response to osmotic stress in plants. Journal of Plant Research, 124 (4):509-525. doi: 10.1007/s10265-011-0412-3 pmid: 21416314
[31]	Gao Lei, Li Hui, Zheng Huan, Tao Jian-min. 2022. Advances in biosynthesis and regulation mechanism of anthocyanins in fruit trees. Jiangsu Journal of Agricultural Sciences, 38 (1):258-267. (in Chinese)
	高磊, 李慧, 郑焕, 陶建敏. 2022. 果树中花色苷的生物合成及其调控机制研究进展. 江苏农业学报, 38 (1):258-267.
[32]	Gao Z, Li Q, Li J, Chen Y, Luo M, Li H, Wang J, Wu Y, Duan S, Wang L, Song S, Xu W, Zhang C, Wang S, Ma C. 2018. Characterization of the ABA receptor VlPYL 1 that regulates anthocyanin accumulation in grape berry skin. Frontiers in Plant Science, 9:592. doi: 10.3389/fpls.2018.00592 URL
[33]	Garcia-Pastor M E, Serrano M, Guillen F, Castillo S, Martinez-Romero D, Valero D, Zapata P J. 2019. Methyl jasmonate effects on table grape ripening,vine yield,berry quality and bioactive compounds depend on applied concentration. Scientia Horticulturae, 247:380-389. doi: 10.1016/j.scienta.2018.12.043
[34]	Gil-Muñoz F, Sánchez-Navarro J A, Besada C, Salvador A, Badenes M L, Naval M D M, Ríos G. 2020. MBW complexes impinge on anthocyanidin reductase gene regulation for proanthocyanidin biosynthesis in persimmon fruit. Scientific Reports, 10 (1):3543. doi: 10.1038/s41598-020-60635-w pmid: 32103143
[35]	Gonzalez A, Zhao M, Leavitt J M F, Lloyd A M. 2008. Regulation of the anthocyanin biosynthetic pathway by the TTG1/bHLH/Myb transcriptional complex in Arabidopsis seedlings. Plant Journal, 53 (5):814-827. doi: 10.1111/tpj.2008.53.issue-5 URL
[36]	Gou J Y, Felippes F F, Liu C J, Weigel D, Wang J W. 2011. Negative regulation of anthocyanin biosynthesis in Arabidopsis by a miR156-targeted SPL transcription factor. Plant Cell, 23 (4):1512-1522. doi: 10.1105/tpc.111.084525 URL
[37]	Gould K S. 2004. Nature's Swiss army knife:the diverse protective roles of anthocyanins in leaves. Journal of Biomedicine and Biotechnology,(5):314-320.
[38]	Gouot J C, Smith J P, Holzapfel B P, Walker A R, Barril C. 2019. Grape berry flavonoids:a review of their biochemical responses to high and extreme high temperatures. Journal of Experimental Botany, 70 (2):397-423. doi: 10.1093/jxb/ery392 URL
[39]	Ha C V, Leyva-González M A, Osakabe Y, Tran U T, Nishiyama R, Watanabe Y, Tanaka M, Seki M, Yamaguchi S, Dong N V, Yamaguchi-Shinozaki K, Shinozaki K, Herrera-Estrella L, Tran L S. 2014. Positive regulatory role of strigolactone in plant responses to drought and salt stress. Proceedings of the National Academy of Sciences, 111 (2):851-856.
[40]	Hauvermale A L, Ariizumi T, Steber C M. 2014. The roles of the GA receptors GID1a,GID1b,and GID1c in sly1-independent GA signaling. Plant Signaling & Behavior, 9 (2):2125-2139.
[41]	He J, Giusti M M. 2010. Anthocyanins:natural colorants with health-promoting properties. Annual Review of Food Science and Technology, 1 (1):163-187. doi: 10.1146/food.2010.1.issue-1 URL
[42]	He Wei, Li Hui-ping, Gao Ping, Han Qin, Liu Bo, Jiang Dong-ze, Lu Si-yu, Wang Li-na. 2022. Effect of exogenous gibberellin treatment on relieving hypocotyl dormancy of Sichuan Paeonia rubra seeds. Agricultural Science & Technology, 23 (2):20-23. (in Chinese)
	何伟, 李会萍, 高平, 韩沁, 刘波, 江东泽, 陆思羽, 王丽娜. 2022. 外源赤霉素处理对解除川赤芍种子下胚轴休眠的影响. 农业科学与技术, 23 (2):20-23.
[43]	Hu B, Li J, Wang D, Wang H, Qin Y, Hu G, Zhao J. 2018. Transcriptome profiling of Litchi chinensis pericarp in response to exogenous cytokinins and abscisic acid. Plant Growth Regulation, 84 (3):437-450. doi: 10.1007/s10725-017-0351-7 URL
[44]	Ito S, Nozoye T, Sasaki E, Imai M, Shiwa Y, Shibata-Hatta M, Ishige T, Fukui K, Ito K, Nakanishi H, Nishizawa N K, Yajima S, Asami T. 2015. Strigolactone regulates anthocyanin accumulation,acid phosphatases production and plant growth under low phosphate condition in Arabidopsis. PLoS ONE, 10 (3):e0119724. doi: 10.1371/journal.pone.0119724 URL
[45]	Jia H, Xie Z, Wang C, Shangguan L, Qian N, Cui M, Liu Z, Zheng T, Wang M, Fang J. 2017. Abscisic acid,sucrose,and auxin coordinately regulate berry ripening process of the Fujiminori grape. Functional & Integrative Genomics, 17 (4):441-457.
[46]	Jiang G, Li Z, Song Y, Zhu H, Lin S, Huang R, Jiang Y, Duan X. 2019. LcNAC 13 physically interacts with LcR1MYB1 to coregulate anthocyanin biosynthesis-related genes during litchi fruit ripening. Biomolecules, 9 (4):135. doi: 10.3390/biom9040135 URL
[47]	Jiu S A O, Guan L, Leng X, Zhang K, Haider M S, Yu X, Zhu X, Zheng T, Ge M, Wang C, Jia H, Shangguan L, Zhang C, Tang X, Abdullah M, Javed H U, Han J, Dong Z, Fang J. 2021. The role of VvMYBA2r and VvMYBA2w alleles of the MYBA2 locus in the regulation of anthocyanin biosynthesis for molecular breeding of grape(Vitis spp.) skin coloration. Plant Biotechnology Journal, 19 (6):1216-1239. doi: 10.1111/pbi.v19.6 URL
[48]	Ju Y L, Liu B C, Xu X L, Wu J R, Sun W, Fang Y L. 2022. Targeted metabolomic and transcript level analysis reveals the effects of exogenous strigolactone and methyl jasmonate on grape quality. Scientia Horticulturae, 299:111009. doi: 10.1016/j.scienta.2022.111009 URL
[49]	Kanzaki S, Ichihi A, Tanaka Y, Fujishige S, Koeda S, Shimizu K. 2020. The R2R3-MYB transcription factor MiMYB 1 regulates light dependent red coloration of‘Irwin’mango fruit skin. Scientia Horticulturae, 272:109567. doi: 10.1016/j.scienta.2020.109567 URL
[50]	Kaplan M, Najda A, Klimek K, Borowy A. 2019. Effect of gibberellic acid (GA₃) inflorescence application on content of bioactive compounds and antioxidant potential of grape (Vitis L.)‘Einset Seedless’berries. South African Journal of Enology and Viticulture, 40 (1):1-10.
[51]	Karppinen K, Tegelberg P, Häggman H, Jaakola L. 2018. Abscisic acid regulates anthocyanin biosynthesis and gene expression associated with cell wall modification in ripening bilberry (Vaccinium myrtillus L.) fruits. Frontiers in Plant Science, 9:1259. doi: 10.3389/fpls.2018.01259 pmid: 30210522
[52]	Kieber J J, Schaller G E. 2018. Cytokinin signaling in plant development. Development,doi:10.1242/dev.149344.
[53]	Koes R, Verweij W, Quattrocchio F. 2005. Flavonoids:a colorful model for the regulation and evolution of biochemical pathways. Trends in Plant Science, 10 (5):236-242. doi: 10.1016/j.tplants.2005.03.002 URL
[54]	Kok D, Bal E. 2018. Enhancing skin color and phenolic compounds of cv. Red Globe table grape(V. vinifera L.) utilizing of different preharvest treatments. Erwerbs-Obstbau, 60 (1):75-81. doi: 10.1007/s10341-017-0352-8 URL
[55]	Kour J, Kohli S K, Khanna K, Bakshi P, Sharma P, Singh A D, Ibrahim M, Devi K, Sharma N, Ohri P, Skalicky M, Brestic M, Bhardwaj R, Landi M, Sharma A. 2021. Brassinosteroid signaling,crosstalk and,physiological functions in plants under heavymetal stress. Frontiers in Plant Science, 12:608061. doi: 10.3389/fpls.2021.608061 URL
[56]	Koyama R, Roberto S R, de Souza R T, Borges W F S, Anderson M, Waterhouse A L, Cantu D, Fidelibus M W, Blanco-Ulate B. 2018. Exogenous abscisic acid promotes anthocyanin biosynthesis and increased expression of flavonoid synthesis genes in Vitis vinifera× Vitis labrusca table grapes in a subtropical region. Frontiers in Plant Science, 9:323. doi: 10.3389/fpls.2018.00323 URL
[57]	Lai B, Du L N, Liu R, Hu B, Su W B, Qin Y H, Zhao J T, Wang H C, Hu G B. 2016. Two LcbHLH transcription factors interacting with LcMYB 1 in regulating late structural genes of anthocyanin biosynthesis in Nicotiana and Litchi chinensis during anthocyanin accumulation. Frontiers in Plant Science, 7:166.
[58]	Leyser O. 2018. Auxin signaling. Plant Physiology, 176 (1):465-479. doi: 10.1104/pp.17.00765 pmid: 28818861
[59]	Li D, Luo Z, Mou W, Wang Y, Ying T, Mao L. 2014. ABA and UV-C effects on quality,antioxidant capacity and anthocyanin contents of strawberry fruit (Fragaria ananassa Duch.). Postharvest Biology and Technology, 90:56-62. doi: 10.1016/j.postharvbio.2013.12.006 URL
[60]	Li H, Duan S Y, Sun W L, Wang S F, Zhang J, Song T T, Tian J, Yao Y C. 2022. Identification, through transcriptome analysis of transcription factors that regulate anthocyanin biosynthesis in different parts of red-fleshed apple‘May’fruit. Horticultural Plant Journal, 8 (1):11-21. doi: 10.1016/j.hpj.2021.07.001 URL
[61]	Li L, Li D, Luo Z, Li X. 2016. Proteomic response and quality maintenance in postharvest fruit of strawberry(Fragaria× ananassa) to exogenous cytokinin. Scientific Reports, 6 (1):27094. doi: 10.1038/srep27094
[62]	Li Lixian, Wang Shuo, Chen Ying, Wu Yingtao, Wang Yaqian, Fang Yue, Chen Xuesen, Tian Changping, Feng Shouqian. 2022. PavMYB10.1 promotes anthocyanin accumulation by positively regulating PavRiant in sweet cherry. Acta Horticulturae Sinica, 49 (5):1023-1030. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0732 URL
	李丽仙, 王烁, 陈莹, 邬滢涛, 王雅倩, 房月, 陈学森, 田长平, 冯守千. 2022. 甜樱桃PavMYB10.1促进PavRiant表达和花青苷积累. 园艺学报, 49 (5):1023-1030. doi: 10.16420/j.issn.0513-353x.2021-0732 URL
[63]	Li M, Cheng S, Wang Y, Dong Y. 2020a. Improving fruit coloration,quality attributes,and phenolics content in‘Rainier’and‘Bing’cherries by gibberellic acid combined with homobrassinolide. Journal of Plant Growth Regulation, 39 (3):1130-1139. doi: 10.1007/s00344-019-10049-4
[64]	Li W, Ding Z, Ruan M, Yu X, Peng M, Liu Y. 2017. Kiwifruit R2R3-MYB transcription factors and contribution of the novel AcMYB75 to red kiwifruit anthocyanin biosynthesis. Scientific Reports, 7 (1):1-14. doi: 10.1038/s41598-016-0028-x
[65]	Li W, Nguyen K H, Tran C D, Watanabe Y, Tian C, Yin X, Li K, Yang Y, Guo J, Miao Y, Yamaguchi S, Tran L P. 2020b. Negative roles of strigolactone-related SMXL6,7 and 8 proteins in drought resistance in Arabidopsis. Biomolecules,doi:10.3390/biom10040607.
[66]	Li Wei-lin, Wei Zhi-wen, Yang Hai-yan, Wu Wen-Long. 2022. Research on the mechanism of cytokinins involved in nitrogen regulation of lateral branch formation and development in higher plants. Journal of Shenyang Agricultural University, 53 (3):373-383. (in Chinese)
	李维林, 魏智文, 杨海燕, 吴文龙. 2022. 细胞分裂素参与氮素调控高等植物侧枝形成和发育的作用机制研究. 沈阳农业大学学报, 53 (3):373-383.
[67]	Li X L, Cui H N, Song X F, Sun C Z, Ding Z, Zhu X Y, Liu X F, Yan L Y. 2023. CsIPT1b and CsUGT85A2 delay female corolla opening in cucumber by regulating the content of cytokinins. Horticultural Plant Journal, 9 (4):754-762. doi: 10.1016/j.hpj.2022.09.005 URL
[68]	Liu H, Liu Z, Wu Y, Zheng L, Zhang G. 2021a. Regulatory mechanisms of anthocyanin biosynthesis in apple and pear. International Journal of Molecular Sciences,doi:10.3390/ijms22168441.
[69]	Liu N. 2019. Effects of IAA and ABA on the immature peach fruit development process. Horticultural Plant Journal, 5 (4):145-154. doi: 10.1016/j.hpj.2019.01.005
[70]	Liu R, Lai B, Hu B, Qin Y, Hu G, Zhao J. 2017. Identification of MicroRNAs and their target genes related to the accumulation of anthocyanins in Litchi chinensis by high-throughput sequencing and degradome analysis. Frontiers in Plant Science,doi:10.3389/fpls.2016.02059.
[71]	Liu Xiao-fen, Li Fang, Yin Xue-ren, Xu Chang-jie, Chen Kun-song. 2013. Advances in transcriptional regulation of anthocyanin biosynthesis. Acta Horticulturae Sinica, 40 (11):2295-2306. (in Chinese)
	刘晓芬, 李方, 殷学仁, 徐昌杰, 陈昆松. 2013. 花青苷生物合成转录调控研究进展. 园艺学报, 40 (11):2295-2306.
[72]	Liu Xiao-yue, Wang Rong-hua, Wang Wei-cheng, Liu Na, Liu Xiao-han, Liu Chao-yang, Liu Da-li, Wu Ze-dong, Wang Mao-qian. 2021. Cytokinin concentrations:effect on the germination of sugarbeet seeds. Chinese Agricultural Science Bulletin, 37 (11):9-14. (in Chinese)
	刘小越, 王荣华, 王维成, 刘镎, 刘晓晗, 刘朝阳, 刘大丽, 吴则东, 王茂芊. 2021. 不同浓度细胞分裂素对甜菜种子引发的影响. 中国农学通报, 37 (11):9-14. doi: 10.11924/j.issn.1000-6850.casb2020-0532
[73]	Liu Y, Ma K, Qi Y, Lv G, Ren X, Liu Z, Ma F. 2021b. Transcriptional regulation of anthocyanin synthesis by MYB-bHLH-WDR complexes in kiwifruit (Actinidia chinensis). Journal of Agricultural and Food Chemistry, 69 (12):3677-3691. doi: 10.1021/acs.jafc.0c07037 URL
[74]	Lu Suwen, Zheng Xuanang, Wang Jiayang, Fang Jinggui. 2021. Research progress on the metabolism of flavonoids in grape. Acta Horticulturae Sinica, 48 (12):2506-2524. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0558 URL
	卢素文, 郑暄昂, 王佳洋, 房经贵. 2021. 葡萄类黄酮代谢研究进展. 园艺学报, 48 (12):2506-2524. doi: 10.16420/j.issn.0513-353x.2021-0558 URL
[75]	Lu Yu, Dong Xian-yi, Du Jing-ping, Li Yong-qiang, Wang Ming-lin. 2004. Research progress on anthocyanins. Journal of Shandong Agricultural University (Natural Science Edition),(2):315-320. (in Chinese)
	卢钰, 董现义, 杜景平, 李永强, 王明林. 2004. 花色苷研究进展. 山东农业大学学报 (自然科学版),(2):315-320.
[76]	Luan Li-ying. 2014. Study on the regulation of anthocyanin synthesis of grape and the quality of wine after brassinolide and abscisic acid treatments[Ph. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	栾丽英. 2014. 油菜素内酯和脱落酸对酿酒葡萄花色苷调控及葡萄酒品质影响的研究[博士论文]. 杨凌: 西北农林科技大学.
[77]	Manganaris G A, Vicente A R, Crisosto C H, Labavitch J M. 2008. Effect of delayed storage and continuous ethylene exposure on flesh reddening of ‘Royal Diamond’plums. Journal of the Science of Food and Agriculture, 88 (12):2180-2185. doi: 10.1002/jsfa.v88:12 URL
[78]	Mano Y, Nemoto K. 2012. The pathway of auxin biosynthesis in plants. Journal of experimental Botany, 63 (8):2853-2872. doi: 10.1093/jxb/ers091 pmid: 22447967
[79]	Mhetre V B, Patel V B, Singh S K, Mishra G P, Verma M K, Kumar C, Dahuja A, Kumar S, Singh R, Wasim Siddiqui M. 2022. Unraveling the pathways influencing the berry color and firmness of grapevine cv. Flame Seedless treated with bioregulators using biochemical and RNA-Seq analysis under semi-arid subtropics. Food Chemistry:Molecular Sciences, 5:100116. doi: 10.1016/j.fochms.2022.100116 URL
[80]	Moro L, Hassimotto N M A, Purgatto E. 2017. Postharvest auxin and methyl jasmonate effect on anthocyanin biosynthesis in red raspberry (Rubus idaeus L.). Journal of Plant Growth Regulation, 36 (3):773-782. doi: 10.1007/s00344-017-9682-x URL
[81]	Naing A H, Kim C K. 2018. Roles of R2R3-MYB transcription factors in transcriptional regulation of anthocyanin biosynthesis in horticultural plants. Plant Molecular Biology, 98 (1):1-18. doi: 10.1007/s11103-018-0771-4
[82]	Ni J, Premathilake A T, Gao Y, Yu W, Tao R, Teng Y A O, Bai S A O. 2021. Ethylene-activated PpERF105 induces the expression of the repressor-type R2R3-MYB gene PpMYB140 to inhibit anthocyanin biosynthesis in red pear fruit. Plant Journal, 105 (1):167-181. doi: 10.1111/tpj.v105.1 URL
[83]	Pan Cheng. 2018. Relationship between gene expression of gibberellin biosynthesis and metabolism related enzymes and spring bud dormancy release in tea plant[Ph. D. Dissertation]. Hefei: Anhui Agricultural University. (in Chinese)
	潘铖. 2018. 茶树赤霉素合成和代谢相关酶的基因表达与春芽休眠解除关系研究[博士论文]. 合肥: 安徽农业大学.
[84]	Pan Jia-li, Zhao Yan, Huang Li-hua, Gao Han, Huang Yu, Chen Jin-jun, Zhang Xue-wen. 2020. The polar transport of the auxin in tobacco trichome and its effect on cell elongation. Journal of Hunan Agricultural University (Natural Sciences), 46 (6):686-690. (in Chinese)
	潘嘉立, 赵燕, 黄丽华, 高晗, 黄妤, 陈金军, 张学文. 2020. 生长素在烟草表皮毛中的极性运输及对细胞伸长的影响. 湖南农业大学学报 (自然科学版), 46 (6):686-690.
[85]	Pattyn J, Vaughan-Hirsch J, van de Poel B. 2021. The regulation of ethylene biosynthesis:a complex multilevel control circuitry. New Phytologist, 229 (2):770-782. doi: 10.1111/nph.v229.2 URL
[86]	Peng Y, Thrimawithana A H, Cooney J M, Jensen D J, Espley R V, Allan A C. 2020. The proanthocyanin-related transcription factors MYBC1 and WRKY 44 regulate branch points in the kiwifruit anthocyanin pathway. Scientific Reports, 10 (1):1-19. doi: 10.1038/s41598-019-56847-4
[87]	Peppi M C, Fidelibus M W, Dokoozlian N. 2006. Abscisic acid application timing and concentration affect firmness,pigmentation,and color of‘Flame Seedless’grapes. HortScience, 41 (6):1440-1445. doi: 10.21273/HORTSCI.41.6.1440 URL
[88]	Portu J, López R, Baroja E, Santamaría P, Garde-Cerdán T. 2016. Improvement of grape and wine phenolic content by foliar application to grapevine of three different elicitors:methyl jasmonate,chitosan,and yeast extract. Food Chemistry, 201:213-221. doi: 10.1016/j.foodchem.2016.01.086 URL
[89]	Qi T, Song S, Ren Q, Wu D, Huang H, Chen Y, Fan M, Peng W, Ren C, Xie D. 2011. The jasmonate-ZIM-domain proteins interact with the WD-Repeat/bHLH/MYB complexes to regulate jasmonate-mediated anthocyanin accumulation and trichome initiation in Arabidopsis thaliana. Plant Cell, 23 (5):1795-1814. doi: 10.1105/tpc.111.083261 URL
[90]	Qu S, Wang G, Li M, Yu W, Zhu S. 2022. LcNAC90 transcription factor regulates biosynthesis of anthocyanin in harvested litchi in response to ABA and GA₃. Postharvest Biology and Technology, 194:112109. doi: 10.1016/j.postharvbio.2022.112109 URL
[91]	Rahim M, Busatto N, Trainotti L. 2014. Regulation of anthocyanin biosynthesis in peach fruits. Planta, 240 (5):913-929. doi: 10.1007/s00425-014-2078-2 pmid: 24827911
[92]	Reynolds A, Robbins N, Lee H S, Kotsaki E. 2016. Impacts and interactions of abscisic acid and gibberellic acid on sovereign Coronation and Skookum seedless table grapes. American Journal of Enology and Viticulture, 67 (3):327-338. doi: 10.5344/ajev.2016.15108 URL
[93]	Robson F, Okamoto H, Patrick E, Harris S R, Wasternack C, Brearley C, Turner J G. 2010. Jasmonate and phytochrome A signaling in Arabidopsis wound and shade responses are integrated through JAZ1 stability. Plant Cell, 22 (4):1143-1160. doi: 10.1105/tpc.109.067728 URL
[94]	Ruan J, Zhou Y, Zhou M, Yan J, Khurshid M, Weng W, Cheng J, Zhang K. 2019. Jasmonic acid signaling pathway in plants. International Journal of Molecular Sciences, 20 (10):2479. doi: 10.3390/ijms20102479 URL
[95]	Ruyter-Spira C, Al-Babili S, van der Krol S, Bouwmeester H. 2013. The biology of strigolactones. Trends in Plant Science, 18 (2):72-83. doi: 10.1016/j.tplants.2012.10.003 pmid: 23182342
[96]	Schaart J G, Dubos C, Romero De La Fuente I, van Houwelingen A M M L, de Vos R C H, Jonker H H, Xu W, Routaboul J M, Lepiniec L, Bovy A G. 2013. Identification and characterization of MYB-bHLH-WD40 regulatory complexes controlling proanthocyanidin biosynthesis in strawberry (Fragaria × ananassa) fruits. New Phytologist, 197 (2):454-467. doi: 10.1111/nph.12017 pmid: 23157553
[97]	Schaefer H M, Schaefer V, Levey D J. 2004. How plant-animal interactions signal new insights in communication. Trends in Ecology & Evolution, 19 (11):577-584. doi: 10.1016/j.tree.2004.08.003 URL
[98]	Sheard L B, Tan X, Mao H, Withers J, Ben-Nissan G, Hinds TR, Kobayashi Y, Hsu F F, Sharon M, Browse J, He S Y, Rizo J, Howe G A, Zheng N. 2010. Jasmonate perception by inositol-phosphate-potentiated COI1-JAZ co-receptor. Nature, 468 (7322):400-405. doi: 10.1038/nature09430
[99]	Shen X, Zhao K, Liu L, Zhang K, Yuan H, Liao X, Wang Q, Guo X, Li F, Li T. 2014. A role for PacMYBA in ABA-regulated anthocyanin biosynthesis in red-colored sweet cherry cv. Hong Deng (Prunus avium L.). Plant and Cell Physiology, 55 (5):862-880. doi: 10.1093/pcp/pcu013 URL
[100]	Shi B, Wu H, Zhu W, Zheng B, Wang S, Zhou K, Qian M. 2022. Genome-wide identification and expression analysis of WRKY genes during anthocyanin biosynthesis in the mango (Mangifera indica L.). Agriculture, 12 (6):821. doi: 10.3390/agriculture12060821 URL
[101]	Starkevič P, Paukštytė J, Kazanavičiūtė V, Denkovskienė E, Stanys V, Bendokas V, Šikšnianas T, Ražanskienė A, Ražanskas R. 2015. Expression and anthocyanin biosynthesis-modulating potential of sweet cherry (Prunus avium L.) MYB10 and bHLH genes. PLoS ONE, 10 (5):e0126991. doi: 10.1371/journal.pone.0126991 URL
[102]	Su M, Wang S, Liu W, Yang M, Zhang Z, Wang N, Chen X. 2022. MdJA 2 participates in the brassinosteroid signaling pathway to regulate the synthesis of anthocyanin and proanthocyanidin in red-fleshed apple. Frontiers in Plant Science, 13:830349. doi: 10.3389/fpls.2022.830349 URL
[103]	Su Z, Wang X, Xuan X, Sheng Z, Jia H, Emal N, Liu Z, Zheng T, Wang C, Fang J. 2021. Characterization and action mechanism analysis of VvmiR156b/c/d-VvSPL9 module responding to multiple-hormone signals in the modulation of grape berry color formation. Foods,doi:10.3390/foods10040896.
[104]	Sudheeran P K, Love C, Feygenberg O, Maurer D, Ovadia R, Oren-Shamir M, Alkan N. 2019. Induction of red skin and improvement of fruit quality in‘Kent’,‘Shelly’and‘Maya’mangoes by preharvest spraying of prohydrojasmon at the orchard. Postharvest Biology and Technology, 149:18-26. doi: 10.1016/j.postharvbio.2018.11.014
[105]	Sun J, Wang Y, Chen X, Gong X, Wang N, Ma L, Qiu Y, Wang Y, Feng S. 2017. Effects of methyl jasmonate and abscisic acid on anthocyanin biosynthesis in callus cultures of red-fleshed apple (Malus sieversii f. niedzwetzkyana). Plant Cell,Tissue and Organ Culture, 130 (2):227-237. doi: 10.1007/s11240-017-1217-4 URL
[106]	Sun Jiamao, Cui Quanshi, Wang Yuqing, Si Yajing, Shi Yufan, Bu Haidong, Yuan Hui, Wang Aide. 2022. Effects of brassinolides and methyl jasmonate spraying on the postharvest quality of apple fruit. Acta Horticulturae Sinica, 49 (10):2236-2248. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-0594 URL
	孙嘉茂, 崔全石, 王语晴, 司雅静, 时瑀繁, 卜海东, 袁晖, 王爱德. 2022. 苹果采前喷施EBR与MeJA对采后品质的影响. 园艺学报, 49 (10):2236-2248. doi: 10.16420/j.issn.0513-353x.2022-0594 URL
[107]	Sun Jian-xia, Zhang Yan, Hu Xiao-song, Wu Ji-hong, Liao Xiao-jun. 2009. Structural stability and degradation mechanisms of anthocyanins. Scientia Agricultura Sinica, 42 (3):996-1008. (in Chinese)
	孙建霞, 张燕, 胡小松, 吴继红, 廖小军. 2009. 花色苷的结构稳定性与降解机制研究进展. 中国农业科学, 42 (3):996-1008.
[108]	Sun W J, Ji X L, Song L Q, Wang X F, You C X, Hao Y J. 2021. Functional identification of MdSMXL8.2,the homologous gene of strigolactones pathway repressor protein gene in Malus × domestica. Horticultural Plant Journal, 7 (4):275-285. doi: 10.1016/j.hpj.2021.01.001 URL
[109]	Tanaka Y, Sasaki N, Fau-Ohmiya A. 2008. Biosynthesis of plant pigments:anthocyanins,betalains and carotenoids. Plant Journal, 54 (4):733-749. doi: 10.1111/tpj.2008.54.issue-4 URL
[110]	Tang T, Zhou H, Wang L, Zhao J, Ma L, Ling J, Li G, Huang W, Li P, Zhang Y. 2022. Post-harvest application of methyl jasmonate or prohydrojasmon affects color development and anthocyanins biosynthesis in peach by regulation of sucrose metabolism. Frontiers in Nutrition, 9:871467. doi: 10.3389/fnut.2022.871467 URL
[111]	Thines B, Katsir L, Melotto M, Niu Y, Mandaokar A, Liu G, Nomura K, He S Y, Howe G A, Browse J. 2007. JAZ repressor proteins are targets of the SCFCOI 1 complex during jasmonate signalling. Nature, 448 (7154):661-665. doi: 10.1038/nature05960
[112]	Tian F, Han C, Chen X, Wu X, Mi J, Wan X, Liu Q, He F, Chen L, Yang H, Zhong Y, Qian Z, Zhang F. 2022. PscCYP716A1-mediated brassinolide biosynthesis increases cadmium tolerance and enrichment in poplar. Frontiers in Plant Science,doi:10.3389/fpls.2022.919682.
[113]	Tsurunaga Y, Takahashi T, Katsube T, Kudo A, Kuramitsu O, Ishiwata M, Matsumoto S. 2013. Effects of UV-B irradiation on the levels of anthocyanin,rutin and radical scavenging activity of buckwheat sprouts. Food Chemistry, 141 (1):552-556. doi: 10.1016/j.foodchem.2013.03.032 pmid: 23768393
[114]	Tyagi K, Maoz I, Lapidot O, Kochanek B, Butnaro Y, Shlisel M, Lerno L, Ebeler S E, Lichter A. 2022. Effects of gibberellin and cytokinin on phenolic and volatile composition of Sangiovese grapes. Scientia Horticulturae, 295:110860. doi: 10.1016/j.scienta.2021.110860 URL
[115]	Villarreal N M, Bustamante C A, Civello P M, Martínez G A. 2010. Effect of ethylene and 1-MCP treatments on strawberry fruit ripening. Journal of the Science of Food and Agriculture, 90 (4):683-689. doi: 10.1002/jsfa.3868 pmid: 20355099
[116]	Wang H, Huang H, Huang X. 2007. Differential effects of abscisic acid and ethylene on the fruit maturation of Litchi chinensis Sonn. Plant Growth Regulation, 52 (3):189-198. doi: 10.1007/s10725-007-9189-8 URL
[117]	Wang L, Wang B, Yu H, Guo H, Lin T, Kou L, Wang A, Shao N, Ma H, Xiong G, Li X, Yang J, Chu J, Li J. 2020. Transcriptional regulation of strigolactone signalling in Arabidopsis. Nature, 583 (7815):277-281. doi: 10.1038/s41586-020-2382-x
[118]	Wang P, Ge M, Yu A, Song W, Fang J, Leng X. 2022a. Effects of ethylene on berry ripening and anthocyanin accumulation of‘Fujiminori’grape in protected cultivation. Journal of the Science of Food and Agriculture, 102 (3):1124-1136. doi: 10.1002/jsfa.v102.3 URL
[119]	Wang S, Li L X, Fang Y, Li D, Mao Z, Zhu Z, Chen X S, Feng S Q. 2022b. MdERF1B-MdMYC2 module integrate ethylene and jasmonic acid to regulate the biosynthesis of anthocyanin in apple. Horticulture Research,doi:10.1093/hr/uhac142.
[120]	Wang Y C, Wang N, Xu H F, Jiang S H, Fang H C, Su M Y, Zhang Z Y, Zhang T L, Chen X S. 2018b. Auxin regulates anthocyanin biosynthesis through the Aux/IAA-ARF signaling pathway in apple. Horticulture Research, 5:59. doi: 10.1038/s41438-018-0068-4
[121]	Wang Y, Liu W, Jiang H, Mao Z, Wang N, Jiang S, Xu H, Yang G, Zhang Z, Chen X. 2019. The R2R3-MYB transcription factor MdMYB24-like is involved in methyl jasmonate-induced anthocyanin biosynthesis in apple. Plant Physiology and Biochemistry, 139:273-282. doi: S0981-9428(19)30121-4 pmid: 30925437
[122]	Wang Y, Mao Z, Jiang H, Zhang Z, Wang N, Chen X. 2021. Brassinolide inhibits flavonoid biosynthesis and red-flesh coloration via the MdBEH2.2-MdMYB 60 complex in apple. Journal of Experimental Botany, 72 (18):6382-6399. doi: 10.1093/jxb/erab284 URL
[123]	Wang Y, Sun J, Wang N, Xu H, Qu C, Jiang S, Fang H, Su M, Zhang Z, Chen X. 2018a. MdMYBL2 helps regulate cytokinin-induced anthocyanin biosynthesis in red-fleshed apple (Malus sieversii f. niedzwetzkyana) callus. Functional Plant Biology, 46 (2):187-196. doi: 10.1071/FP17216 URL
[124]	Wasternack C, Hause B. 2019. The missing link in jasmonic acid biosynthesis. Nature Plants, 5 (8):776-777. doi: 10.1038/s41477-019-0492-y pmid: 31358962
[125]	Wei Ding-yi, Gao Shao-fan, Su Hai-ying, Xie Ling-li, Xu Ben-bo. 2022. Effects of gibberellin and its inhibitors on lodging resistance of rape. Jiangsu Agricultural Sciences, 50 (12):69-74. (in Chinese)
	韦丁一, 高少凡, 苏海英, 谢伶俐, 许本波. 2022. 赤霉素及其抑制剂对油菜抗倒伏性的影响. 江苏农业科学, 50 (12):69-74.
[126]	Wu T, Liu H T, Zhao G P, Song J X, Wang X L, Yang C Q, Zhai R, Wang Z G, Ma F W, Xu L F. 2020. Jasmonate and ethylene-regulated ethylene response factor 22 promotes lanolin-induced anthocyanin biosynthesis in‘Zaosu’pear (Pyrus bretschneideri Rehd.) fruit. Biomolecules, 10 (2):278. doi: 10.3390/biom10020278 URL
[127]	Xie S, Liu Y, Chen H, Yang B, Ge M, Zhang Z. 2022. Effects of gibberellin applications before flowering on the phenotype,ripening,and flavonoid compounds of Syrah grape berries. Journal of the Science of Food and Agriculture, 102 (13):6100-6111. doi: 10.1002/jsfa.v102.13 URL
[128]	Xie Y, Tan H, Ma Z, Huang J. 2016. DELLA proteins promote anthocyanin biosynthesis via sequestering MYBL2 and JAZ suppressors of the MYB/bHLH/WD 40 complex in Arabidopsis thaliana. Molecular Plant, 9 (5):711-721. doi: S1674-2052(16)00033-2 pmid: 26854848
[129]	Yang Bo, Wei Jia, Li Kunfeng, Wang Chengliang, Ni Junbei, Teng Yuanwen, Bai Songling. 2022. PpyERF060-PpyABF3-PpyMADS71regulates ethylene signaling pathway- mediated pear bud dormancy process. Acta Horticulturae Sinica, 49 (10):2249-2262. (in Chinese)
	杨博, 魏佳, 李坤峰, 王程亮, 倪隽蓓, 滕元文, 白松龄. 2022. PpyERF060-PpyABF3-PpyMADS71调控乙烯信号通路介导的梨芽休眠进程. 园艺学报, 49 (10):2249-2262. doi: 10.16420/j.issn.0513-353x.2022-0585 URL
[130]	Yang C Q, Sha G Y, Wei T, Ma B Q, Li C Y, Li P M, Zou Y J, Xu L F, Ma F W. 2021. Linkage map and QTL mapping of red flesh locus in apple using a R1R1 × R6R6 population. Horticultural Plant Journal, 7 (5):393-400. doi: 10.1016/j.hpj.2020.12.008 URL
[131]	Yang F W, Feng X Q. 2015. Abscisic acid biosynthesis and catabolism and their regulation roles in fruit ripening. Phyton, 84 (2):444-453. doi: 10.32604/phyton.2015.84.444 URL
[132]	Yang Hai-yan, Yan Zhi-xiang, Fan Su-fan, Wu Wen-long, Lü Lian-fei, Li Wei-lin. 2020. Effects of different gibberellic acid concentrations on the fruits quality of blueberry cultivar‘Anna’. Northern Horticulture,(23):39-43. (in Chinese)
	杨海燕, 严志祥, 樊苏帆, 吴文龙, 闾连飞, 李维林. 2020. 不同浓度赤霉素处理对基质栽培蓝莓‘安娜’果实品质的影响. 北方园艺,(23):39-43.
[133]	Yang Weihai, Zeng Lizhen, Xiao Qiusheng, Shi Shengyou. 2021. Changes of fruit abscission and carbohydrate,ABA and related genes expression in the pericarp and fruit abscission zone of longan under starvation stress. Acta Horticulturae Sinica, 48 (8):1457-1469. (in Chinese)
	杨为海, 曾利珍, 肖秋生, 石胜友. 2021. 饥饿胁迫下龙眼落果与果皮和离区糖、ABA及相关基因表达的变化. 园艺学报, 48 (8):1457-1469. doi: 10.16420/j.issn.0513-353x.2020-0538 URL
[134]	Yao G, Ming M, Allan AC, Gu C, Li L, Wu X, Wang R, Chang Y, Qi K, Zhang S, Wu J. 2017. Map-based cloning of the pear gene MYB 114 identifies an interaction with other transcription factors to coordinately regulate fruit anthocyanin biosynthesis. Plant Journal, 92 (3):437-451. doi: 10.1111/tpj.2017.92.issue-3 URL
[135]	Yuan Qiao-hui. 2019. Preliminary study on the regulation of cytokinin on the quality of strawberry fruit[M. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	原乔慧. 2019. 细胞分裂素对草莓果实品质调控作用初探[硕士论文]. 南京: 南京农业大学.
[136]	Zaharah S S, Singh Z, Symons G M, Reid J B. 2012. Role of brassinosteroids,ethylene,abscisic acid,and indole-3-acetic acid in mango fruit ripening. Journal of Plant Growth Regulation, 31 (3):363-372. doi: 10.1007/s00344-011-9245-5 URL
[137]	Zeng Tuo, Li Jiawen, Zhou Li, Li Jinjin, Shi Anqi, Fu Hansen, Luo Jing, Zheng Riru, Wang Yuanyuan, Wang Caiyun. 2021. Advances in the mutualistic and antagonistic interactions between flower colors and the pollinators of ornamental plants. Acta Horticulturae Sinica, 48 (10):2001-2017. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0523 URL
	曾拓, 李伽文, 周黎, 李进进, 史安琪, 付瀚森, 罗靖, 郑日如, 王媛媛, 王彩云. 2021. 观赏植物花色与授粉昆虫相互适应关系的研究进展. 园艺学报, 48 (10):2001-2017. doi: 10.16420/j.issn.0513-353x.2021-0523 URL
[138]	Zhang J, Xu H, Wang N, Jiang S, Fang H, Zhang Z, Yang G, Wang Y, Su M, Xu L, Chen X. 2018. The ethylene response factor MdERF1B regulates anthocyanin and proanthocyanidin biosynthesis in apple. Plant Molecular Biology, 98 (3):205-218. doi: 10.1007/s11103-018-0770-5 pmid: 30182194
[139]	Zhang L, Song C, Guo D, Guo L, Hou X, Wang H. 2022a. Identification of differentially expressed miRNAs and their target genes in response to brassinolide treatment on flowering of tree peony (Paeonia ostii). Plant Signaling & Behavior, 17 (1):2056364.
[140]	Zhang Y, Liu Z, Liu J, Lin S, Wang J, Lin W, Xu W. 2017. GA-DELLA pathway is involved in regulation of nitrogen deficiency-induced anthocyanin accumulation. Plant Cell Reports, 36 (4):557-569. doi: 10.1007/s00299-017-2102-7 pmid: 28275852
[141]	Zhang Y, Xu Y, Huang D, Xing W, Wu B, Wei Q, Xu Y, Zhan R, Ma F, Song S. 2022b. Research progress on the MYB transcription factors in tropical fruit. Tropical Plants, 1 (1):1-15.
[142]	Zhang Z, Shi Y, Ma Y, Yang X, Yin X, Zhang Y, Xiao Y, Liu W, Li Y, Li S, Liu X, Grierson D, Allan AC, Jiang G, Chen K. 2020. The strawberry transcription factor FaRAV1 positively regulates anthocyanin accumulation by activation of FaMYB10 and anthocyanin pathway genes. Plant Biotechnology Journal, 18 (11):2267-2279. doi: 10.1111/pbi.v18.11 URL
[143]	Zhang Zhe, Liu Fang, Zhang Chao, Song Shui-shan. 2014. Advances in research on strigolactones and its signaling pathway. Journal of Biology, 31 (2):64-68. (in Chinese)
	张哲, 刘方, 张超, 宋水山. 2014. 独脚金素内酯及其信号转导通路的研究进展. 生物学杂志, 31 (2):64-68.
[144]	Zhao M, Li J, Zhu L, Chang P, Li L, Zhang L. 2019. Identification and characterization of MYB-bHLH-WD 40 regulatory complex members controlling anthocyanidin biosynthesis in blueberry fruits development. Genes, 10 (7):496. doi: 10.3390/genes10070496 URL
[145]	Zheng Xiaodong, Xi Xiangli, Li Yuqi, Sun Zhijuan, Ma Changqing, Han Mingsan, Li Shaoxuan, Tian Yike, Wang Caihong. 2022. Effects and regulating mechanism of exogenous brassinosteroids on the growth of Malus hupehensis under saline-alkali stress. Acta Horticulturae Sinica, 49 (7):1404-1414. (in Chinese)
	郑晓东, 袭祥利, 李玉琪, 孙志娟, 马长青, 韩明三, 李少旋, 田义轲, 王彩虹. 2022. 油菜素内酯对盐碱胁迫下平邑甜茶幼苗生长的影响及调控机理研究. 园艺学报, 49 (7):1401-1414. doi: 10.16420/j.issn.0513-353x.2021-0499 URL
[146]	Zhou H, Kui L W, Wang H, Gu C, Dare A P, Espley R V, He H, Allan A C, Han Y. 2015. Molecular genetics of blood-fleshed peach reveals activation of anthocyanin biosynthesis by NAC transcription factors. Plant Journal, 82 (1):105-121. doi: 10.1111/tpj.2015.82.issue-1 URL
[147]	Zimmermann I M, Heim M A, Weisshaar B l, Uhrig J F. 2004. Comprehensive identification of Arabidopsis thaliana MYB transcription factors interacting with R/B-like BHLH proteins. Plant Journal, 40 (1):22-34. doi: 10.1111/j.1365-313X.2004.02183.x pmid: 15361138
[148]	Zwack P J, Rashotte A M. 2013. Cytokinin inhibition of leaf senescence. Plant Signaling & Behavior, 8 (7):e24737.

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

激素信号调控果树果实花色苷合成机制研究进展

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

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激素信号调控果树果实花色苷合成机制研究进展

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

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