Ahmad, M., Lin, C., Cashmore, A.R., 1995. Mutations throughout an Arabidopsis blue-light photoreceptor impair blue-light-responsive anthocyanin accumulation and inhibition of hypocotyl elongation. The Plant Journal 8 (5), 653-658.

An, J.P., Qu, F.J., Yao, J.F., Wang, X.N., You, C.X., Wang, X.F., Hao, Y.J., 2017. The bZIP transcription factor MdHY5 regulates anthocyanin accumulation and nitrate assimilation in apple. Hortic. Res.-England 4, 17023. https://doi.org/10.1038/hortres.2017.23.

Ang, L.H., Chattopadhyay, S., Wei, N., Oyama, T., Okada, K., Batschauer, A., Deng, X.W., 1998. Molecular interaction between COP1 and HY5 defines a regulatory switch for light control of Arabidopsis development. Mol. Cell 1 (2), 213-222. https://doi.org/10.1016/s1097-2765(00)80022-2.

Bai, S., Tao, R., Tang, Y., Yin, L., Ma, Y., Ni, J., Yan, X., Yang, Q., Wu, Z., Zeng, Y., Teng, Y., 2019a. BBX16, a B-Box protein, positively regulates light-induced anthocyanin accumulation by activating MYB10 in red pear. Plant Biotechnol. J. 17 (10), 1985-1997. https://doi.org/10.1111/pbi.13114.

Bai, S., Tao, R., Yin, L., Ni, J., Yang, Q., Yan, X., Yang, F., Guo, X., Li, H., Teng, Y., 2019b. Two b-box proteins, PpBBX18 and PpBBX 21, antagonistically regulate anthocyanin biosynthesis via competitive association with Pyrus pyrifolia elongated hypocotyl 5 in the peel of pear fruit. Plant J. 100 (6), 1208-1223. https://doi.org/10.1111/tpj.14510.

Bhagat, P.K., Verma, D., Sharma, D., Sinha, A.K., 2021. HY5 and ABI5 transcription factors physically interact to fine tune light and ABA signaling in Arabidopsis. Plant Mol.Biol. 107 (1-2), 117-127. https://doi.org/10.1007/s11103-021-01187-z.

Bhatnagar, A., Singh, S., Khurana, J.P., Burman, N., 2020. HY5-COP1: the central module of light signaling pathway. J. Plant Biochem. Biotechnol. 29 (4), 590-610.

Chaves, I., Pokorny, R., Byrdin, M., Hoang, N., Ritz, T., Brettel, K., Essen, L., Van Der Horst, G.T., Batschauer, A., Ahmad, M., 2011. The cryptochromes: blue light photoreceptors in plants and animals. Annu. Rev. Plant Biol. 62 (1), 335-364.

Chen, Z., An, Y., Wang, L., 2023. Ala reverses aba-induced stomatal closure by modulating PP2AC and SnRK2. 6 activity in apple leaves. Hortic. Res.-England 10 (6), d67.

Corrales, M., Toepfl, S., Butz, P., Knorr, D., Tauscher, B., 2008. Extraction of anthocyanins from grape by-products assisted by ultrasonics, high hydrostatic pressure or pulsed electric fields: a comparison. Innov. Food Sci. Emerg. Technol. 9 (1), 85-91.

Dong, H., Hu, C., Liu, C., Wang, J., Zhou, Y., Yu, J., 2021. Elongated hypocotyl 5 mediates blue light-induced starch degradation in tomato. J. Exp. Bot. 72 (7), 2627-2641. https://doi.org/10.1093/jxb/eraa604.

Fang, P., Yan, M., Chi, C., Wang, M., Zhou, Y., Zhou, J., Shi, K., Xia, X., Foyer, C.H., Yu, J., 2019. Brassinosteroids act as a positive regulator of photoprotection in response to chilling stress. Plant Physiol. 180 (4), 2061-2076. https://doi.org/10.1104/pp.19.00088.

Gangappa, S.N., Crocco, C.D., Johansson, H., Datta, S., Hettiarachchi, C., Holm, M., Botto, J.F., 2013. The arabidopsis b-box protein bbx25 interacts with hy5, negatively regulating BBX22 expression to suppress seedling photomorphogenesis. Plant Cell 25 (4), 1243-1257. https://doi.org/10.1105/tpc.113.109751.

Guo, Z., Xu, J., Wang, Y., Hu, C., Shi, K., Zhou, J., Xia, X., Zhou, Y., Foyer, C.H., Yu, J., 2021. The PHYB-dependent induction of hy5 promotes iron uptake by systemically  activating fer expression. EMBO Rep. 22 (7), e51944. https://doi.org/10.15252/embr.202051944.

Hardtke, C.S., Gohda, K., Osterlund, M.T., Oyama, T., Okada, K., Deng, X.W., 2000. HY5 stability and activity in Arabidopsis is regulated by phosphorylation in its COP1 binding domain. Embo J. 19 (18), 4997-5006. https://doi.org/10.1093/emboj/19.18.4997.

Holm, M., Hardtke, C.S., Gaudet, R., Deng, X.W., 2001. Identification of a structural motif that confers specific interaction with the WD40 repeat domain of Arabidopsis cop1. EMBO J. 20 (1-2), 118-127. https://doi.org/10.1093/emboj/20.1.118.

Jaakola, L., 2013. New insights into the regulation of anthocyanin biosynthesis in fruits. Trends Plant Sci. 18 (9), 477-483. https://doi.org/10.1016/j.tplants.2013.06.003.

Jia, H., Zuo, Q., Sadeghnezhad, E., Zheng, T., Chen, X., Dong, T., Fang, J., 2023. HDAC19 recruits ERF4 to the MYB5a promoter and diminishes anthocyanin accumulation during grape ripening. The Plant Journal 113 (1), 127-144.

Job, N., Lingwan, M., Masakapalli, S.K., Datta, S., 2022. Transcription factors BBX11 and HY5 interdependently regulate the molecular and metabolic responses to UV-B. Plant Physiol. 189 (4), 2467-2480. https://doi.org/10.1093/plphys/kiac195.

Jonassen, E.M., Lea, U.S., Lillo, C., 2008. HY5 and HYH are positive regulators of nitrate reductase in seedlings and rosette stage plants. Planta 227 (3), 559-564. https://doi.org/10.1007/s00425-007-0638-4.

Kami, C., Lorrain, S., Hornitschek, P., Fankhauser, C., 2010. Light-regulated plant growth and development. Curr. Top. Dev. Biol. 91, 29-66. https://doi.org/10.1016/S0070-2153(10)91002-8.

Kim, J.Y., Song, J.T., Seo, H.S., 2017. COP1 regulates plant growth and development in response to light at the post-translational level. J. Exp. Bot. 68 (17), 4737-4748. https://doi.org/10.1093/jxb/erx312.

Lau, O.S., Deng, X.W., 2012. The photomorphogenic repressors COP1 and det1: 20 years later. Trends Plant Sci. 17 (10), 584-593.

Lee, H.S., Wicker, L., 1991. Anthocyanin pigments in the skin of lychee fruit. J. Food Sci. 56 (2), 466-468.

Li, D., Ye, G., Li, J., Lai, Z., Ruan, S., Qi, Q., Wang, Z., Duan, S., Jin, H.L., Wang, H.B., 2023. High light triggers flavonoid and polysaccharide synthesis through DoHY5-dependent signaling in dendrobium officinale. Plant J. 115 (4), 1114-1133. https://doi.org/10.1111/tpj.16284.

Li, Q.F., He, J.X., 2016. BZR1 interacts with HY5 to mediate brassinosteroid- and light-regulated cotyledon opening in Arabidopsis in darkness. Mol. Plant. 9 (1), 113-125. https://doi.org/10.1016/j.molp.2015.08.014.

Li, Y., Mao, K., Zhao, C., Zhao, X., Zhang, R., Zhang, H., Shu, H., Hao, Y., 2013. Molecular cloning and functional analysis of a blue light receptor gene MdCRY2 from apple (Malus domestica). Plant Cell Reports 32, 555-566.

Li, Y., Xu, P., Chen, G., Wu, J., Liu, Z., Lian, H., 2020. FvbHLH9 functions as a positive regulator of anthocyanin biosynthesis by forming a HY5-bHLH9 transcription complex in strawberry fruits. Plant Cell Physiol. 61 (4), 826-837. https://doi.org/10.1093/pcp/pcaa010.

Li, Z., Zhang, C., Guo, Y., Niu, W., Wang, Y., Xu, Y., 2017. Evolution and expression analysis reveal the potential role of the HD-zip gene  family in regulation of embryo abortion in grapes (Vitis vinifera L.). BMC Genomics 18 (1), 744. https://doi.org/10.1186/s12864-017-4110-y.

Lin, C., Ahmad, M., Gordon, D., Cashmore, A.R., 1995. Expression of an Arabidopsis cryptochrome gene in transgenic tobacco results in hypersensitivity to blue, UV-A, and green light. Proceedings of the National Academy of Sciences 92 (18), 8423-8427.

Lin, C., Shalitin, D., 2003. Cryptochrome structure and signal transduction. Annu. Rev. Plant Biol. 54 (1), 469-496.

Lin, R., Ding, L., Casola, C., Ripoll, D.R., Feschotte, C., Wang, H., 2007. Transposase-derived transcription factors regulate light signaling in Arabidopsis. Science 318 (5854), 1302-1305. https://doi.org/10.1126/science.1146281.

Liu, C.C., Ahammed, G.J., Wang, G.T., Xu, C.J., Chen, K.S., Zhou, Y.H., Yu, J.Q., 2018a. Tomato CRY1a plays a critical role in the regulation of phytohormone homeostasis, plant development, and carotenoid metabolism in fruits. Plant, Cell & Environment 41 (2), 354-366.

Liu, C.C., Chi, C., Jin, L.J., Zhu, J., Yu, J.Q., Zhou, Y.H., 2018b. The bZIP transcription factor HY5 mediates CRY1a-induced anthocyanin biosynthesis in tomato. Plant, Cell & Environment 41 (8), 1762-1775.

Liu, W., Zhang, L., Ma, L., Yuan, H., Wang, A., 2023b. The HY5 transcription factor negatively regulates ethylene production by inhibiting ACS1 expression under blue light conditions in pear. Hortic. Plant J. 9 (5), 920-930.

Liu, X., Cheng, X., Cao, J., Zhu, W., Sun, Y., Lin, N.,Li Wan, X., Liu, L., 2023c. UV-b regulates seasonal greening of albino leaves by modulating CsHY5-inhibiting chlorophyll biosynthesis in Camellia sinensis cv. Huangkui. Plant Sci. 328, 111569. https://doi.org/10.1016/j.plantsci.2022.111569.

Liu, Y., Tang, L., Wang, Y., Zhang, L., Xu, S., Wang, X., He, W., Zhang, Y., Lin, Y., Wang, Y., Li, M., Wang, X., Zhang, Y., Luo, Y., Chen, Q., Tang, H., 2023a. The blue light signal transduction module FaCRY1-FaCOP1-FaHY5 regulates anthocyanin accumulation in cultivated strawberry. Front. Plant Sci. 14, 1144273. https://doi.org/10.3389/fpls.2023.1144273.

Livak, K.J., Schmittgen, T.D., 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2 ^−ΔΔCT Method. Methods 25 (4), 402-408.

Maxwell, B.B., Andersson, C.R., Poole, D.S., Kay, S.A., Chory, J., 2003. HY5, Circadian Clock-Associated 1, and a cis-element, DET1 dark response element, mediate DET1 regulation of chlorophyll a/b-binding protein 2 expression. Plant Physiol. 133 (4), 1565-1577.

Ni, J., Liao, Y., Zhang, M., Pan, C., Yang, Q., Bai, S., Teng, Y., 2022. Blue light simultaneously induces peel anthocyanin biosynthesis and flesh carotenoid/sucrose biosynthesis in mango fruit. J. Agric. Food Chem. 70 (50), 16021-16035. https://doi.org/10.1021/acs.jafc.2c07137.

Osterlund, M.T., Hardtke, C.S., Wei, N., Deng, X.W., 2000. Targeted destabilization of HY5 during light-regulated development of Arabidopsis. Nature 405 (6785), 462-466.

Podolec, R., Ulm, R., 2018. Photoreceptor-mediated regulation of the COP1/SPA E3 ubiquitin ligase. Curr. Opin. Plant Biol. 45, 18-25.

Qiu, Z., Wang, H., Li, D., Yu, B., Hui, Q., Yan, S., Huang, Z., Cui, X., Cao, B., 2019. Identification of candidate HY5-dependent and -independent regulators of anthocyanin biosynthesis in tomato. Plant Cell Physiol. 60 (3), 643-656. https://doi.org/10.1093/pcp/pcy236.

Samkumar, A., Jones, D., Karppinen, K., Dare, A.P., Sipari, N., Espley, R.V., Martinussen, I., Jaakola, L., 2021. Red and blue light treatments of ripening bilberry fruits reveal differences in signalling through abscisic acid-regulated anthocyanin biosynthesis. Plant Cell Environ. 44 (10), 3227-3245. https://doi.org/10.1111/pce.14158.

Seo, H.S., Yang, J., Ishikawa, M., Bolle, C., Ballesteros, M.L., Chua, N., 2003. LAF1 ubiquitination by COP1 controls photomorphogenesis and is stimulated by SPA1. Nature 423 (6943), 995-999.

Shao, D., Zhu, Q.H., Liang, Q., Wang, X., Li, Y., Sun, Y., Zhang, X., Liu, F., Xue, F., Sun, J., 2022. Transcriptome analysis reveals differences in anthocyanin accumulation in cotton (Gossypium hirsutum L.) Induced by red and blue light. Front. Plant Sci. 13, 788828. https://doi.org/10.3389/fpls.2022.788828.

Shi, L., Cao, S., Chen, W., Yang, Z., 2014. Blue light induced anthocyanin accumulation and expression of associated genes in chinese bayberry fruit. Sci. Hortic. 179, 98-102.

Shin, D.H., Choi, M., Kim, K., Bang, G., Cho, M., Choi, S.B., Choi, G., Park, Y.I., 2013. Hy5 regulates anthocyanin biosynthesis by inducing the transcriptional activation of the MYB75/PAP1 transcription factor in Arabidopsis. FEBS Lett. 587 (10), 1543-1547. https://doi.org/10.1016/j.febslet.2013.03.037.

Stracke, R., Favory, J.J., Gruber, H., Bartelniewoehner, L., Bartels, S., Binkert, M., Funk, M., Weisshaar, B., Ulm, R., 2010. The Arabidopsis bzip transcription factor HY5 regulates expression of the PFG1/MYB12 gene in response to light and ultraviolet-b radiation. Plant, cell & environment 33 (1), 88-103.

Tamura, K., Stecher, G., Peterson, D., Filipski, A., Kumar, S., 2013. MEGA6: molecular evolutionary genetics analysis version 6.0. Mol. Biol. Evol. 30 (12), 2725-2729. https://doi.org/10.1093/molbev/mst197.

Tao, R., Bai, S., Ni, J., Yang, Q., Zhao, Y., Teng, Y., 2018. The blue light signal transduction pathway is involved in anthocyanin accumulation in 'Red zaosu' pear. Planta 248 (1), 37-48. https://doi.org/10.1007/s00425-018-2877-y.

Wang, F., Zhang, L., Chen, X., Wu, X., Xiang, X., Zhou, J., Xia, X., Shi, K., Yu, J., Foyer, C.H., Zhou, Y., 2019. SlHY5 integrates temperature, light, and hormone signaling to balance plant  growth and cold tolerance. Plant Physiol. 179 (2), 749-760. https://doi.org/10.1104/pp.18.01140.

Wang, H., Ma, L., Li, J., Zhao, H., Deng, X.W., 2001. Direct interaction of Arabidopsis cryptochromes with COP1 in light control development. Science 294 (5540), 154-158.

Wang, X., Wang, Q., Nguyen, P., Lin, C., 2014. Cryptochrome-mediated light responses in plants. Enzymes 35, 167-189. https://doi.org/10.1016/B978-0-12-801922-1.00007-5.

Wang, Y., Zhang, X., Zhao, Y., Yang, J., He, Y., Li, G., Ma, W., Huang, X., Su, J., 2020. Transcription factor PyHY5 binds to the promoters of pywd40 and pymyb10 and regulates its expression in red pear ‘Yunhongli no. 1’. Plant Physiol. Biochem. 154, 665-674.

Xiao, Y., Chu, L., Zhang, Y., Bian, Y., Xiao, J., Xu, D., 2021. HY5: a pivotal regulator of light-dependent development in higher plants. Front. Plant Sci. 12, 800989. https://doi.org/10.3389/fpls.2021.800989.

Xing, Y., Sun, W., Sun, Y., Li, J., Zhang, J., Wu, T., Song, T., Yao, Y., Tian, J., 2023. MPK6-mediated HY5 phosphorylation regulates light-induced anthocyanin accumulation in apple fruit. Plant Biotechnol. J. 21 (2), 283-301. https://doi.org/10.1111/pbi.13941.

Xiong, H., Lu, D., Li, Z., Wu, J., Ning, X., Lin, W., Bai, Z., Zheng, C., Sun, Y., Chi, W., Zhang, L., Xu, X., 2023. The DELLA-ABI4-HY5 module integrates light and gibberellin signals to regulate hypocotyl elongation. Plant Commun. 4 (5), 100597.

Yan, J., Liu, J., Yang, S., Jiang, C., Liu, Y., Zhang, N., Sun, X., Zhang, Y., Zhu, K., Peng, Y., Bu, X., Wang, X., Ahammed, G.J., Meng, S., Tan, C., Liu, Y., Sun, Z., Qi, M., Wang, F., Li, T., 2023a. Light quality regulates plant biomass and fruit quality through a  photoreceptor-dependent HY5-LHC/CYCB module in tomato. Hortic. Res.-England 10 (12), d219.

Yan, Y., Zhao, J., Lin, S., Li, M., Liu, J., Raymond, O., Vergne, P., Kong, W., Wu, Q., Zhang, X., Bao, M., Bendahmane, M., Fu, X., 2023b. Light-mediated anthocyanin biosynthesis in rose petals involves a balanced  regulatory module comprising transcription factors RhHY5, RhMYB114a, and RhMYB3b. J. Exp. Bot. 74 (18), 5783-5804.

Yang, C., Li, F., Ji, X., Wang, J., 2011. An effective method for rna extraction from grapevine berry skins. African Journal of Biotechnology 10 (45), 9032-9035.

Yang, H., Tang, R., Cashmore, A.R., 2001. The signaling mechanism of Arabidopsis CRY1 involves direct interaction with COP1. The Plant Cell 13 (12), 2573-2587.

Yang, J., Lin, R., Sullivan, J., Hoecker, U., Liu, B., Xu, L., Deng, X.W., Wang, H., 2005. Light regulates COP1-mediated degradation of hfr1, a transcription factor essential for light signaling in Arabidopsis. The Plant Cell 17 (3), 804-821.

Yeh, C.S., Wang, Z., Miao, F., Ma, H., Kao, C.T., Hsu, T.S., Yu, J.H., Hung, E.T., Lin, C.C., Kuan, C.Y., Tsai, N.C., Zhou, C., Qu, G.Z., Jiang, J., Liu, G., Wang, J.P., Li, W., Chiang, V.L., Chang, T.H., Lin, Y.J., 2019. A novel synthetic-genetic-array-based yeast one-hybrid system for high discovery rate and short processing time. Genome Res. 29 (8), 1343-1351.

Zhang, C., Wu, Y., Liu, X., Zhang, J., Li, X., Lin, L., Yin, R., 2022. Pivotal roles of elongated hypocotyl5 in regulation of plant development and fruit metabolism in tomato. Plant Physiol. 189 (2), 527-540.

Zhang, H., He, H., Wang, X., Wang, X., Yang, X., Li, L., Deng, X.W., 2011. Genome‐wide mapping of the hy5-mediated genenetworks in arabidopsis that involve both transcriptional and post‐transcriptional regulation. The Plant Journal 65 (3), 346-358.

Zhang, Y., Butelli, E., Martin, C., 2014. Engineering anthocyanin biosynthesis in plants. Curr. Opin. Plant Biol. 19, 81-90.

Zhang, Y., Jiang, L., Li, Y., Chen, Q., Ye, Y., Zhang, Y., Luo, Y., Sun, B., Wang, X., Tang, H., 2018. Effect of red and blue light on anthocyanin accumulation and differential gene expression in strawberry (Fragaria x ananassa). Molecules 23 (4).

Zheng, C., Ma, J.Q., Ma, C.L., Shen, S.Y., Liu, Y.F., Chen, L., 2019. Regulation of growth and flavonoid formation of tea plants (Camellia sinensis) by blue and green light. J. Agric. Food Chem. 67 (8), 2408-2419.

Zoratti, L., Karppinen, K., Luengo Escobar, A., Häggman, H., Jaakola, L., 2014. Light-controlled flavonoid biosynthesis in fruits. Front. Plant Sci. 5, 534.