果实成熟进程中蛋白质翻译后修饰的功能与作用机制

doi:10.16420/j.issn.0513-353x.2026-0158

摘要/Abstract

摘要：

果实成熟的复杂生物学过程受多层次分子网络精密调控，蛋白质翻译后修饰（post-translational modifications，PTM）是其核心调控节点。本文中系统梳理了磷酸化、泛素化、乙酰化、糖基化等经典PTM，以及甲基化、巴豆酰化、巯基化、氧化还原修饰、S-亚硝基化等新兴PTM在果实成熟中的功能与作用机制。各类PTM通过调控乙烯、脱落酸（ABA）等激素信号，耦合MAPK、CDPK、SnRK2等激酶级联与组蛋白乙酰化/甲基化等表观遗传修饰，实现对呼吸跃变型与非跃变型果实成熟的差异化调控；PTM间还存在磷酸化—泛素化耦联等串扰机制，协同调控转录因子，代谢酶的活性、稳定性与亚细胞定位，进而介导色素合成、细胞壁降解、糖酸代谢、挥发性香气物质合成及营养物质积累，最终影响果实色泽、质地、风味、香气和营养品质。此外，低温、光照等环境因子及乙烯、NO、H₂S等外源信号可介导PTM动态重塑，精细调节果实成熟、品质形成与采后维持。深入解析PTM的调控网络与分子机制，既能完善果实成熟的分子调控理论体系，也为基于PTM靶点的基因编辑育种、采后精准调控策略提供理论支撑。

关键词: 果实, 成熟, 品质, 翻译后修饰, 外源信号介导, 采后调控

Abstract:

Fruit ripening is a complex biological process that is precisely regulated by multi-level molecular networks，in which protein post-translational modifications（PTMs）act as central regulatory hubs. This review systematically summarizes the functions and mechanisms of classical PTMs，including phosphorylation，ubiquitination，acetylation and glycosylation，as well as emerging PTMs such as methylation，crotonylation，persulfidation，redox-related modifications and S-nitrosylation during fruit ripening. Different PTMs differentially regulate the ripening of climacteric and non-climacteric fruits by modulating hormone signaling pathways，such as ethylene and abscisic acid（ABA），as well as kinase cascades including MAPK，CDPK and SnRK2，and epigenetic modifications such as histone acetylation and methylation. Crosstalk mechanisms，such as phosphorylation-ubiquitination coupling，also occur among PTMs，coordinately regulating the activity，stability and subcellular localization of transcription factors and metabolic enzymes. These processes further mediate pigment biosynthesis，cell wall degradation，sugar-acid metabolism，volatile compound synthesis and nutrient accumulation，ultimately determining fruit color，texture，taste，aroma and nutritional quality. In addition，environmental factors such as low temperature and light，as well as exogenous signals including ethylene，nitric oxide（NO）and hydrogen sulfide（H₂S），can dynamically remodel PTMs to finely regulate fruit ripening，quality formation and postharvest preservation. A deeper understanding of the regulatory networks and molecular mechanisms of PTMs will not only improve the theoretical framework of fruit ripening regulation，but also provide a foundation for PTM-targeted gene editing breeding and precision postharvest regulation strategies，thereby promoting fruit quality improvement and extending storage life.

Key words: fruit, ripening, quality, post-translational modifications, exogenous signal mediation, postharvest regulation

段程远, 卫玮, 单伟, 陈建业, 陆旺金, 邝健飞, 杨滢滢. 果实成熟进程中蛋白质翻译后修饰的功能与作用机制[J]. 园艺学报, 2026, 53(7): 1853-1870.

DUAN Chengyuan, WEI Wei, SHAN Wei, CHEN Jianye, LU Wangjin, KUANG Jianfei, YANG Yingying. Functions and Mechanisms of Protein Post-Translational Modifications During Fruit Ripening[J]. Acta Horticulturae Sinica, 2026, 53(7): 1853-1870.

图/表 2

图1 果实成熟多级激酶磷酸化信号调控机制模式图乙烯、ABA、Ca2+、蔗糖及发育和环境因素等上游信号经MAPK级联及CDPK、SnRK2等通路传递，通过调控物种特异性下游靶标活性，最终影响乙烯生物合成、细胞壁降解、色素积累以及果实成熟和采后货架期等生理过程

Fig. 1 Schematic diagram of the multi-step kinase phosphorylation signaling regulation mechanism in fruit ripening Upstream signals such as ethylene，ABA，Ca2+，sucrose，and developmental and environmental factors are transmitted via the MAPK cascade and pathways involving CDPK and SnRK2. By regulating the activity of species-specific downstream targets，these signals ultimately influence physiological processes such as ethylene biosynthesis，cell wall degradation，pigment accumulation，fruit ripening，and postharvest shelf life.（Wang et al.，2014；Yoon，2015；Wu et al.，2021；Yang et al.，2022；Li et al.，2023；Wei et al.，2023；Zhou et al.，2024a；Jia et al.，2025；Tan et al.，2025）

图2 多重翻译后修饰（PTM）协同调控果实成熟与品质形成的分子机制内源激素和外源环境信号通过磷酸化、泛素化、乙酰化、糖基化及表观遗传修饰等途径协同调控基因表达、蛋白稳定性和代谢重塑，最终影响果实成熟进程、品质形成及采后性状

Fig. 2 Molecular mechanism of multiple post-translational modifications（PTMs）coordinately regulating fruit ripening and quality formation Endogenous hormones and exogenous environmental signals work in concert to regulate gene expression，protein stability，and metabolic remodeling through pathways such as phosphorylation，ubiquitination，acetylation，glycosylation，and epigenetic modifications，ultimately influencing fruit ripening，quality development，and postharvest traits（Meli et al.，2010；Komander & Rape，2012；Wang et al.，2014；Yoon，2015；Han et al.，2016；Wu et al.，2019，2021，2023；Corpas et al.，2021；Pattyn et al.，2021；Ming et al.，2023；Sougrakpam et al.，2023；An et al.，2024；Ma et al.，2024；Zhou et al.，2024b；Jia et al.，2025；Li et al.，2025b；Tan et al.，2025；Zhao et al.，2025）

参考文献 94

[45]	Ma X, Jia Q X, Li S, Chen Z T, Ming X, Zhao Y, Zhou D X. 2023. An enhanced network of energy metabolism,lysine acetylation,and growth-promoting protein accumulation is associated with heterosis in elite hybrid rice. Plant Communications, 4 (4):100560. doi: 10.1016/j.xplc.2023.100560 URL
[46]	Meli V S, Ghosh S, Prabha T N, Chakraborty N, Chakraborty S, Datta A. 2010. Enhancement of fruit shelf life by suppressing N-glycan processing enzymes. Proceedings of the National Academy of Sciences of the United States of America, 107 (6):2413-2418.
[47]	Millar A H, Heazlewood J L, Giglione C, Holdsworth M J, Bachmair A, Schulze W X. 2019. The scope,functions,and dynamics of posttranslational protein modifications. Annual Review of Plant Biology,70:119-151.
[48]	Ming Y, Jiang L, Ji D. 2023. Epigenetic regulation in tomato fruit ripening. Frontiers in Plant Science, 14:1269090. doi: 10.3389/fpls.2023.1269090 URL
[49]	Mu L J, Wang X K, Ma Y X, Zhao A J, Han S B, Li R, Lei K, Ji L S, Li P. 2024. Apple glycosyltransferase MdUGT73AR4 glycosylates ABA to regulate stomatal movement involved in drought stress. International Journal of Molecular Sciences, 25 (11):5672. doi: 10.3390/ijms25115672 URL
[50]	Naik J, Rajput R, Singh S, Stracke R, Pandey A. 2025. Heat-responsive MaHSF 11 transcriptional activator positively regulates flavonol biosynthesis and flavonoid B-ring hydroxylation in banana(Musa acuminata). The Plant Journal, 121 (5):e70084. doi: 10.1111/tpj.v121.5 URL
[51]	Nareshkumar V, Harish Chandar S R, D’Orso F, Aparna J, Girija Sangari M, Sumega R, Viswanathan C, Appunu C. 2026. Genome editing for trait optimization:the role of CRISPR,base editing and prime editing in modern era of plant science. Journal of Plant Biochemistry and Biotechnology, 35 (1):119-144. doi: 10.1007/s13562-025-01005-3
[1]	An C T, Liu Z S, Pan X J, Li A L, Wei W Y, Yang T Y, Liao W B, Wang C L. 2024. Effect of histone modifications on fruit ripening. Physiologia Plantarum, 176 (6):e14639. doi: 10.1111/ppl.v176.6 URL
[2]	Anzalone A V, Randolph P B, Davis J R, Sousa A A, Koblan L W, Levy J M, Chen P J, Christopher W, Newby G A, Aditya R, Raguram A, Liu D R. 2019. Search-and-replace genome editing without double-strand breaks or donor DNA. Nature, 576 (7785):149-157. doi: 10.1038/s41586-019-1711-4
[3]	Baldwin A, Lechon T, Marchbank A, Scofield S, Lieu K, Wilson C L, Ludlow R A, Herbert R. J, Nützmann H W, Rogers H J. 2025. The H3K27me 3 histone mark correlates with repression of colour and aroma development post-harvest in strawberry fruit. Journal of Experimental Botany, 76 (9):2487-2499. doi: 10.1093/jxb/erae464 URL
[4]	Bao L F, Wang W Y, Li M Y, Liu J, Liu J H, Alifu G, Wang D S, Liang X Q, Mao T Y, Zhai Y L. 2025. Reactive oxygen species-post translational modifications-central carbon metabolism regulatory loop:coordination of redox homeostasis and carbon flux allocation in plants under abiotic stress. Frontiers in Plant Science, 16:1637328. doi: 10.3389/fpls.2025.1637328 URL
[5]	Bennett J, Meiyalaghan S, Nguyen H M, Boldingh H, Cooney J, Elborough C, Araujo L D, Barrell P, Lin W K, Plunkett B J, Martin D, Espley R V. 2023. Exogenous abscisic acid and sugar induce a cascade of ripening events associated with anthocyanin accumulation in cultured Pinot Noir grape berries. Frontiers in Plant Science, 14:1324675. doi: 10.3389/fpls.2023.1324675 URL
[52]	Pan Q W, Guo S P, Ding J, Zhou Y, Huang X R, Qi Q, Wang F, Wu H, Li Y, Gu T T. 2024. Dynamic histone modification signatures coordinate developmental programs in strawberry fruit ripening. Horticulture Research, 11 (8):uhae158.
[53]	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
[54]	Pickart C M. 2001. Mechanisms underlying ubiquitination. Annual Review of Biochemistry,70:503-533.
[55]	Roundtree I A, Evans M E, Pan T, He C. 2017. Dynamic RNA modifications in gene expression regulation. Cell, 169 (7):1187-1200. doi: S0092-8674(17)30638-4 pmid: 28622506
[56]	Ruan Banjun, Dai Peng, Wang Wei, Sun Jianbin, Zhang Wentao, Yan Zhen, Yang Jinghua. 2014. Progress on post-translational modification of proteins. Chinese Journal of Cell Biology,(7):1027-1037. (in Chinese)
	阮班军, 代鹏, 王伟, 孙建斌, 张文涛, 颜真, 杨静华. 2014. 蛋白质翻译后修饰研究进展. 中国细胞生物学学报,(7):1027-1037.
[57]	Sonawane P D, Gharat S A, Jozwiak A, Barbole R, Heinicke S, Almekias-Siegl E, Meir S, Rogachev I, Connor S E O, Giri A P. 2023. A BAHD-type acyltransferase concludes the biosynthetic pathway of non-bitter glycoalkaloids in ripe tomato fruit. Nature Communications, 14 (1):4540. doi: 10.1038/s41467-023-40092-5
[58]	Song C, Zhou R L, Zhang H H, Zhang J Y, Gao H Y, Wang R R, Cui Y J, Liu X Y, Wang Z Y, Song S W, Jiao J, Wang M M, Wan R, Shi J L, Hao P B, Zhang K X, Bai T H, Zheng X B. 2026. Histone deacetylase inhibitor affects apple wax and aroma metabolism. Journal of Agricultural and Food Chemistry, 74 (15):3545-3557.
[6]	Cai L, Sutter B M, Li B, Tu B P. 2011. Acetyl-CoA induces cell growth and proliferation by promoting the acetylation of histones at growth genes. Molecular Cell,42:426-437.
[7]	Chen T H, Wei W, Shan W, Kuang J F, Chen J Y, Lu W J, Yang Y Y. 2023. MaHDA6-MaNAC 154 module regulates the transcription of cell wall modification genes during banana fruit ripening. Postharvest Biology and Technology, 198:112237. doi: 10.1016/j.postharvbio.2022.112237 URL
[8]	Choudhary C, Kumar C, Gnad F, Nielsen M L, Rehman M, Walther T C, Olsen J V, Mann M. 2009. Lysine acetylation targets protein complexes and co-regulates major cellular functions. Science, 325 (5942):834-840. doi: 10.1126/science.1175371 pmid: 19608861
[9]	Chung C R, Tang Y, Chiu Y P, Li S, Hsieh W K, Yao L, Chiang Y C, Pang Y, Chen G T, Chou K C, Paik Y S, Tran P L, Lin C P, Kao Y M, Chen Y J, Chang W C, Hsu J B K, Horng J T, Lee T Y. 2024. dbPTM 2025 update:comprehensive integration of PTMs and proteomic data for advanced insights into cancer research. Nucleic Acids Research, 53 (D1):D377-D386.
[10]	Contreras-de La Rosa P A, Aragón-Rodríguez C, Ceja-López J A, García-Arteaga K F, De-la-Peña C. 2022. Lysine crotonylation:a challenging new player in the epigenetic regulation of plants. Journal of Proteomics, 255:104488. doi: 10.1016/j.jprot.2022.104488 URL
[59]	Sougrakpam Y, Babuta P, Deswal R. 2023. Nitric oxide(NO)modulates low temperature-stress signaling via S-nitrosation,a NO PTM,inducing ethylene biosynthesis inhibition leading to enhanced post-harvest shelf-life of agricultural produce. Physiology and Molecular Biology of Plants, 29 (12):2051-2065. doi: 10.1007/s12298-023-01371-z pmid: 38222283
[60]	Sun C, Yao G F, Li L X, Zhao Y Q, Hu K D, Zhang C H, Zhang H. 2023. E3 ligase BRG3 persulfidation delays tomato ripening by reducing ubiquitination of the repressor WRKY71. Plant Physiology, 192 (1):616-632. doi: 10.1093/plphys/kiad070 pmid: 36732924
[61]	Sun L, Fan X, Zhang Y, Jiang J F, Sun Hai S, Liu Chong H. 2016. Transcriptome analysis of genes involved in anthocyanins biosynthesis and transport in berries of black and white spine grapes(Vitis davidii). Hereditas, 153:17. doi: 10.1186/s41065-016-0021-1 URL
[62]	Sun Y F, Ji K, Liang B, Du Y W, Jiang L, Wang J, Kai W B, Zhang Y S, Zhai X W, Chen P, Wang H Q, Leng P. 2017. Suppressing ABA uridine diphosphate glucosyltransferase(SlUGT75C1)alters fruit ripening and the stress response in tomato. The Plant Journal, 91 (4):574-589. doi: 10.1111/tpj.2017.91.issue-4 URL
[63]	Tan J, Zhou Z, Feng H, Feng H Q, Zhang J T, Zhang R K, Chen Z K, Niu Y J, Liu F Y, Deng Z P. 2025. Proteome and phosphoproteome of tomato fruit identify REDUCED CHLOROPLAST COVERAGE 1a as a ripening regulator. Genomics,Proteomics & Bioinformatics,https://doi.org/10.1093/gpbjnl/qzaf050.
[64]	Tian Shiping. 2013. Molecular regulatory mechanisms of fruit ripening and senescence. Chinese Bulletin of Botany, 48 (5):481-488. (in Chinese) doi: 10.3724/SP.J.1259.2013.00481 URL
[11]	Corpas F J, González-gordo S, Muñoz-vargas M A, Rodríguez-Ruiz M, Palma J M. 2021. The modus operandi of hydrogen sulfide(H₂S)-dependent protein persulfidation in higher plants. Antioxidants, 10 (11):1686. doi: 10.3390/antiox10111686 URL
[12]	Cui X K, Wang J X, Li K, Lv B S, Hou B Ki, Ding Z J. 2024. Protein post-translational modifications in auxin signaling. Journal of Genetics and Genomics, 51 (3):279-291. doi: 10.1016/j.jgg.2023.07.002 URL
[13]	de Bruyn F, Maertens J, Beauprez J, Soetaert W, de Mey M. 2015. Biotechnological advances in UDP-sugar based glycosylation of small molecules. Biotechnology Advances, 33 (2):288-302. doi: 10.1016/j.biotechadv.2015.02.005 pmid: 25698505
[14]	Guo J E. 2022. Histone deacetylase gene SlHDT 1 regulates tomato fruit ripening by affecting carotenoid accumulation and ethylene biosynthesis. Plant Science, 318:111235. doi: 10.1016/j.plantsci.2022.111235 URL
[15]	Guo J E, Hu Z L, Li F F, Zhang L C, Yu X H, Tang B Y, Chen G P. 2017a. Silencing of histone deacetylase SlHDT 3 delays fruit ripening and suppresses carotenoid accumulation in tomato. Plant Science:An International Journal of Experimental Plant Biology,265:29-38.
[16]	Guo J E, Hu Z L, Zhu M K, Li F F, Zhu Z G, Lu Y, Chen G P. 2017b. The tomato histone deacetylase SlHDA 1 contributes to the repression of fruit ripening and carotenoid accumulation. Scientific Reports, 7 (1):7930. doi: 10.1038/s41598-017-08512-x
[17]	Han Y C, Kuang J F, Chen J Y, Liu X C, Xiao Y Y, Fu C C, Wang J N, Wu K Q, Lu W J. 2016. Banana transcription factor maerf 11 recruits histone deacetylase mahda1 and represses the expression of MaACO1 and expansins during fruit ripening. Plant Physiology, 171 (2):1070-1084.
[64]	田世平. 2013. 果实成熟和衰老的分子调控机制. 植物学报, 48 (5):481-488. doi: 10.3724/SP.J.1259.2013.00481
[65]	Tikunov Y M, Molthoff J, de Vos R C H, Beekwilder J, van Houwelingen A, van der Hooft J J J, Nijenhuis-de Vries M, Labrie C W, Verkerke W, van de Geest H, Viquez Zamora M, Presa S, Rambla J L, Granell A, Hall R D, Bovy A G. 2013. Non-smoky glycosyltransferase 1 prevents the release of smoky aroma from tomato fruit. The Plant Cell, 25 (8):3067-3078. doi: 10.1105/tpc.113.114231 pmid: 23956261
[66]	Wang N, Liu W J, Mei Z X, Zhang S H, Zou Q, Yu L, Jiang S H, Fang H C, Zhang Z Y, Chen Z J, Wu S J, Cheng L L, Chen X S. 2024. A functional InDel in the WRKY 10 promoter controls the degree of flesh red pigmentation in apple. Advanced Science, 11 (30):2400998. doi: 10.1002/advs.v11.30 URL
[67]	Wang W, Wang Y, Chen T, Qin G Z, Tian S P. 2022. Current insights into posttranscriptional regulation of fleshy fruit ripening. Plant Physiology, 192 (3):1785-1798. doi: 10.1093/plphys/kiac483 URL
[68]	Wang Y Y, Kong L X, Wang W H, Qin G Z. 2023. Global ubiquitinome analysis reveals the role of E3 ubiquitin ligase FaBRIZ in strawberry fruit ripening. Journal of Experimental Botany, 74 (1):214-232. doi: 10.1093/jxb/erac400 URL
[69]	Wang Y Y, Wang W H, Cai J H, Zhang Y R, Qin G Z, Tian S P. 2014. Tomato nuclear proteome reveals the involvement of specific E2 ubiquitin-conjugating enzymes in fruit ripening. Genome Biology, 15 (12):548. doi: 10.1186/s13059-014-0548-2 URL
[18]	Hershko A, Ciechanover A. 1998. The ubiquitin system. Annual Review of Biochemistry,67:425-479.
[19]	Hu Y N, Han Z Y, Wang T, Li H, Li Q Q, Wang S, Tian J, Wang Y, Zhang X Z, Xu X F, Han Z H, Lü P T, Wu T. 2022. Ethylene response factor MdERF4 and histone deacetylase MdHDA 19 suppress apple fruit ripening through histone deacetylation of ripening-related genes. Plant Physiology, 188 (4):2166-2181. doi: 10.1093/plphys/kiac016 URL
[20]	Huang F L, Sun M M, Yao Z J, Zhou J, Bai Q, Chen X X, Huang Y, Shen Y Y. 2024. Protein kinase SnRK2.6 phosphorylates transcription factor bHLH 3 to regulate anthocyanin homeostasis during strawberry fruit ripening. Journal of Experimental Botany, 75 (18):5627-5640. doi: 10.1093/jxb/erae250 URL
[21]	Hunter T. 1995. Protein kinases and phosphatases: The Yin and Yang of protein phosphorylation and signaling. Cell, 80 (2):225-236. doi: 10.1016/0092-8674(95)90405-0 pmid: 7834742
[22]	Irfan M, Kumar P, Kumar V, Datta A. 2022. Fruit ripening specific expression of β-D-N-acetylhexosaminidase(β-Hex)gene in tomato is transcriptionally regulated by ethylene response factor SlERF.E4. Plant Science, 323:111380. doi: 10.1016/j.plantsci.2022.111380 URL
[23]	Jia H R, Shi Y, Dai Z R, Sun Y F, Shu X, Li B J, Wu R, Lv S Z, Shou J, Yang X F, Jiang G H, Zhang Y C, Allan A C, Chen K S. 2025. Phosphorylation of the strawberry MADS-box CMB1 regulates ripening via the catabolism of abscisic acid. New Phytologist, 246 (4):1627-1646. doi: 10.1111/nph.70065 pmid: 40172024
[70]	Wei W, Luo Q, Yang Y Y, Wu C J, Kuang J F, Chen J Y, Lu W J, Shan W. 2024. E3 ubiquitin ligase MaRZF1 modulates high temperature-induced green ripening of banana by degrading MaSGR1. Plant,Cell & Environment, 47 (4):1128-1140. doi: 10.1111/pce.v47.4 URL
[71]	Wei Y, Jin J T, Xu Y X, Liu W T, Yang G X, Bu H D, Li T, Wang A. 2021. Ethylene-activated MdPUB24 mediates ubiquitination of MdBEL 7 to promote chlorophyll degradation in apple fruit. The Plant Journal, 108 (1):169-182. doi: 10.1111/tpj.v108.1 URL
[72]	Wei Y, Liu Z, Lv T X, Xu Y X, Wei Y J, Liu W T, Liu L, Wang A D, Li T. 2023. Ethylene enhances MdMAPK3-mediated phosphorylation of MdNAC 72 to promote apple fruit softening. The Plant Cell, 35 (8):2887-2909. doi: 10.1093/plcell/koad122 URL
[73]	Wu B P, Cao X M, Liu H R, Zhu C Q, Klee H, Zhang B, Chen K S. 2019. UDP-glucosyltransferase PpUGT85A 2 controls volatile glycosylation in peach. Journal of Experimental Botany, 70 (3):925-936. doi: 10.1093/jxb/ery419 URL
[74]	Wu C J, Deng W, Shan W, Liu X C, Zhu L S, Cai D ling, Wei W, Yang Y Y, Chen J Y, Lu W J, Kuang J F. 2023. Banana MKK1 modulates fruit ripening via the MKK1-MPK6-3/11-4-bZIP21 module. Cell Reports, 42 (8):112832. doi: 10.1016/j.celrep.2023.112832 URL
[24]	Jia H F, Zuo Q Q, Sadeghnezhad E, Zheng T, Chen X Q, Dong T Y, Fang J G. 2023a. HDAC19 recruits ERF4 to the MYB5a promoter and diminishes anthocyanin accumulation during grape ripening. The Plant Journal:For Cell and Molecular Biology, 113 (1):127-144. doi: 10.1111/tpj.v113.1 URL
[25]	Jia W, Liu G S, Zhang P Y, Li H li, Peng Z Z, Wang Y X, Jemrić T, Fu D Q. 2023b. The ubiquitin-26S proteasome pathway and its role in the ripening of fleshy fruits. International Journal of Molecular Sciences, 24 (3):2750. doi: 10.3390/ijms24032750 URL
[26]	Kaulfürst-soboll H, Mertens-beer M, Brehler R, Albert M, von Schaewen A. 2021. Complex N-glycans are important for normal fruit ripening and seed development in tomato. Frontiers in Plant Science, 12:635962. doi: 10.3389/fpls.2021.635962 URL
[27]	Komander D, Rape M. 2012. The ubiquitin code. Annual Review of Biochemistry,81:203-229.
[28]	Kytidou K, Artola M, Overkleeft H S, Aerts J M F G. 2020. Plant glycosides and glycosidases:a treasure-trove for therapeutics. Frontiers in Plant Science, 11:357. doi: 10.3389/fpls.2020.00357 URL
[29]	Leszczuk A, Kalaitzis P, Blazakis K N, Zdunek A. 2020. The role of arabinogalactan proteins(AGPs)in fruit ripening-a review. Horticulture Research, 7 (1):176. doi: 10.1038/s41438-020-00397-8 pmid: 33328442
[30]	L’Hernault S W, Rosenbaum J L. 1985. Chlamydomonas alpha-tubulin is posttranslationally modified by acetylation on the epsilon-amino group of a lysine. Biochemistry, 24 (2):473-478. doi: 10.1021/bi00323a034 pmid: 3919761
[31]	Li G M, Feng B H, Zhu Q H, Jiang K F, Zhang T. 2026. Protein post-translational modifications in plant abiotic stress responses. Plants, 15 (1):52. doi: 10.3390/plants15010052 URL
[32]	Li M Y, Wang L G, Liu Y T, Lin Y X, Zhang Y T, Long Y, Luo C Y, Zhang Y, Chen Q, Chen P W, Wang Y, Wang X R, Tang H R, Luo Y. 2022a. Characterization and regulation mechanism analysis of ubiquitin-conjugating family genes in strawberry reveals a potential role in fruit ripening. BMC Plant Biology, 22 (1):39. doi: 10.1186/s12870-021-03421-8
[33]	Li Q Q, Wang T, Xu C, Li M S, Tian J, Wang Y, Zhang X Z, Xu X F, Han Z H, Wu T. 2022b. MdMADS 6 recruits histone deacetylase MdHDA19 to repress the expression of the carotenoid synthesis-related gene MdCCD1 during fruit ripening. Plants, 11 (5):668. doi: 10.3390/plants11050668 URL
[34]	Li T, Liu L, Yang G X, Cai Y C, Wang Y D, Sun B W, Sun L, Liu W T, Wang A D. 2025a. Ethylene‐activated E 3 ubiquitin ligase MdEAEL1 promotes apple fruit softening by facilitating the dissociation of transcriptional repressor complexes. Advanced Science, 12 (22):2417393. doi: 10.1002/advs.v12.22 URL
[35]	Li T, Zeng J, Yang X Q, Garcia-Caparros P, Duan X W. 2024. The role of protein post-translational modifications in fruit ripening. Horticulturae, 10 (10):1042. doi: 10.3390/horticulturae10101042 URL
[36]	Li Y Y, Meng F L, You H W, Shao Z Y, Xu C Y, Li Y N, Li S W, Liu L H, Wang Q M. 2025b. SlWRKY14 integrates carotenoid and flavonoid biosynthetic pathways to regulate coloration and quality of tomato fruits. Cell Reports, 44 (10):116369. doi: 10.1016/j.celrep.2025.116369 URL
[37]	Li X J, Martín-Pizarro C, Zhou L L, Hou B Z, Wang Y Y, Shen Y Y, Li B B, Posé D, Qin G Z. 2023. Deciphering the regulatory network of the NAC transcription factor FvRIF,a key regulator of strawberry(Fragaria vesca)fruit ripening. The Plant Cell, 35 (11):4020-4045. doi: 10.1093/plcell/koad210 URL
[38]	Li X J, Li Q, Qin G Z, Li B B. 2025c. Protein post-translational modifications:key switches coordinating fruit ripening regulatory networks. Horticulture Research, 13 (3):uhaf351.
[39]	Liu Jing, Li Yachao, Zhou Mengyan, Wu Peng, Feima Xiangqing, Li Ming. 2021. Advances in plant protein post-translational modification omics. Biotechnology Bulletin, 37 (1):67-76. (in Chinese) doi: 10.13560/j.cnki.biotech.bull.1985.2020-0946
	刘静, 李亚超, 周梦岩, 吴鹏飞, 马祥庆, 李明. 2021. 植物蛋白质翻译后修饰组学研究进展. 生物技术通报, 37 (1):67-76. doi: 10.13560/j.cnki.biotech.bull.1985.2020-0946
[40]	Liu K D, Yuan C C, Li H L, Chen K Y, Lu L S, Shen C, Zheng X L. 2018. A qualitative proteome-wide lysine crotonylation profiling of papaya(Carica papaya L.). Scientific Reports, 8 (1):8230. doi: 10.1038/s41598-018-26676-y
[41]	Liu Y D. 2023. Phosphoproteomic analysis highlights phosphorylation of primary metabolism-related proteins in response to postharvest cold storage and subsequent shelf-life in tomato fruit. Postharvest Biology and Technology, 205:112486. doi: 10.1016/j.postharvbio.2023.112486 URL
[42]	Liu Z, Yan J P, Li D K, Luo Q, Yan Q, Liu Z B, Ye L M, Wang J M, Li X F, Yang Y. 2015. UDP-glucosyltransferase71c5,a major glucosyltransferase,mediates abscisic acid homeostasis in Arabidopsis. Plant Physiology, 167 (4):1659-1670. doi: 10.1104/pp.15.00053 URL
[43]	Luo Q, Wei W, Yang Y Y, Wu C J, Chen J Y, Lu W J, Kuang J F, Shan W. 2023. E3 ligase MaNIP1 degradation of NON-YELLOW COLORING1 at high temperature inhibits banana degreening. Plant Physiology, 192 (3):1969-1981. doi: 10.1093/plphys/kiad096 pmid: 36794407
[75]	Wu C J, Shan W, Liu X C, Zhu L S, Wei W, Yang Y Y, Guo Y F, Bouzayen M, Chen J Y, Lu W J, Kuang J F. 2021. Phosphorylation of transcription factor bZIP21 by MAP kinase MPK6-3 enhances banana fruit ripening. Plant Physiology, 188 (3):1665-1685. doi: 10.1093/plphys/kiab539 URL
[76]	Xie Q, Tian Y L, Hu Z L, Zhang L C, Tang B Y, Wang Y S, Li J, Chen G P. 2021. Novel translational and phosphorylation modification regulation mechanisms of tomato(Solanum lycopersicum)fruit ripening revealed by integrative proteomics and phosphoproteomics. International Journal of Molecular Sciences, 22 (21):11782. doi: 10.3390/ijms222111782 URL
[77]	Xie W S, Xiao Y Y, Wei W, Kuang J F, Lu W J, Chen J Y, Yang Y Y. 2026. Methionine oxidation-regulated MaERF95L controls starch and sucrose metabolism in postharvest banana during ripening. Journal of Integrative Plant Biology, 68 (2):439-454. doi: 10.1111/jipb.70075
[78]	Xu J H, Xiong L, Yao J L, Zhao P L, Jiang S H, Sun X H, Dong C H, Jiang H Y, Xu X Y, Zhang Y G. 2024. Hypermethylation in the promoter regions of flavonoid pathway genes is associated with skin color fading during‘Daihong’apple fruit development. Horticulture Research, 11 (3):uhae031.
[79]	Xu Q, Yue Y P, Liu B, Chen Z T, Ma X, Wang J, Zhao Y, Zhou D X. 2023. ACL and HAT1 form a nuclear module to acetylate histone H4K5 and promote cell proliferation. Nature Communications, 14 (1):3265. doi: 10.1038/s41467-023-39101-4 pmid: 37277331
[44]	Ma L L, Zuo J H, Bai C M, Fu A Z, Wang Q, Zhou Z J, Deng Z P, Tan J J, Bouzayen M, Zheng Y Y. 2024. The dynamic N1-methyladenosine RNA methylation provides insights into the tomato fruit ripening. The Plant Journal, 120 (5):2014-2030. doi: 10.1111/tpj.v120.5 URL
[80]	Xu Y X. 2022. Effects of calcium application on apple fruit softening during storage revealed by proteomics and phosphoproteomics. Horticultural Plant Journal, 8 (4):408-422. doi: 10.1016/j.hpj.2022.03.002 URL
[81]	Yan J G, Shan S S, Li J K, Zhang Z K, Yang J L, Zhang W L, Song H M, Xu X B, Duan W H. 2025. DNA methylation profile changes in CpG islands of ethylene-signaling genes regulated by melatonin were involved in alleviating chilling injury of postharvest tomato fruit. International Journal of Molecular Sciences, 26 (13):6170. doi: 10.3390/ijms26136170 URL
[82]	Yan H L, Jiang G X, Wu F W, Li Z W, Xiao L, Jiang Y M, Duan X W. 2021. Sulfoxidation regulation of transcription factor NAC 42 influences its functions in relation to stress-induced fruit ripening in banana. Journal of Experimental Botany, 72 (2):682-699. doi: 10.1093/jxb/eraa474 URL
[83]	Yang M, Lin Y X, Zhang Y T, Li M Y, Chen Q, Zhang Y, Tang H R, Luo Y. 2025. Acetylated FaGAPC2 modulates citrate metabolism in strawberry fruit and participates in tapetum degradation to affect pollen development. Scientia Horticulturae, 352:114436. doi: 10.1016/j.scienta.2025.114436 URL
[84]	Yang Y Y, Wu C J, Shan W, Wei W, Zhao Y ting, Kuang J F, Chen J Y, Jiang Y M, Lu W J. 2022. Mitogen-activated protein kinase 14-mediated phosphorylation of MaMYB4 negatively regulates banana fruit ripening. Horticulture Research, 10 (1):uhac243.
[85]	Yoon G M. 2015. New insights into the protein turnover regulation in ethylene biosynthesis. Molecules and Cells, 38 (7):597-603. doi: 10.14348/molcells.2015.0152 pmid: 26095506
[86]	Yu Y, Jiang K. 2026. Metabolic mediators at the Nexus:How SAM,Acetyl-CoA,and NAD⁺ bridge phytohormone signaling and epigenetic regulation. Current Opinion in Plant Biology, 89:102832. doi: 10.1016/j.pbi.2025.102832 URL
[87]	Zhang Li, Fan Weijuan, Zheng Zhenying, Yang Jun, Kang Le, Wang Hongxia. 2023. Current research progress on glycosylation,methylation and acylation modifications of anthocyanins. Molecular Plant Breeding, 21 (7):2378-2387. (in Chinese)
	张丽, 范维娟, 郑臻颖, 杨俊, 康乐, 王红霞. 2023. 花青素糖基化、甲基化和酰基化修饰的研究现状. 分子植物育种, 21 (7):2378-2387.
[88]	Zhang M, Tan F Q, Fan Y J, Wang T T, Song X, Xie K D, Wu X M, Zhang F, Deng X X, Grosser J W, Guo W W. 2022. Acetylome reprograming participates in the establishment of fruit metabolism during polyploidization in citrus. Plant Physiology, 190 (4):2519-2538. doi: 10.1093/plphys/kiac442 pmid: 36135821
[89]	Zhang Y, Zeng L. 2020. Crosstalk between ubiquitination and other post-translational protein modifications in plant immunity. Plant Communications, 1 (4):100041. doi: 10.1016/j.xplc.2020.100041 URL
[90]	Zhao X Y, Ou R H, Cui T H, Liao J C, Mei Y Y, Wu Y D, Wang Q Y, Yang Y, Liu Y L, Liu Y Q, Xu C J, Pan Y J, Gong Q Q, Deng Z P, Zhang B, Li X, Chen K S. 2025. Quantitative glycoproteomics identifies target N-glycoproteins of the Golgi-class I α-mannosidase SlMNSI1 that is important for tomato fruit development. Plant Communications, 6 (12):101537. doi: 10.1016/j.xplc.2025.101537 URL
[91]	Zhou X, Lei Z, An P. 2024a. Post-translational modification of WRKY transcription factors. Plants, 13 (15):2040. doi: 10.3390/plants13152040 URL
[92]	Zhou Y J, Li Z W, Su X G, Hou H Y, Jiang Y M, Duan X W, Qu H X, Jiang G X. 2024b. Histone deacetylase SlHDA7 impacts fruit ripening and shelf life in tomato. Horticulture Research, 11 (11):uhae234.
[93]	Zhu L S, Chen L, Wu C J, Shan W, Cai D L, Lin Z X, Wei W, Chen J Y, Lu W J, Kuang J F. 2023. Methionine oxidation and reduction of the ethylene signaling component MaEIL9 are involved in banana fruit ripening. Journal of Integrative Plant Biology, 65 (1):150-166. doi: 10.1111/jipb.13363
[94]	Zuo D D, Yang L, Li R Y, Xing J L, Zhang J, Zhang K Y, Liu H N, Pei M S, Wei T L, Guo D L. 2025. Histone H3K 4 demethyltransferase VvJMJ14 promotes H₂O₂ inducing fruit ripening in grape. The Plant Journal, 124 (5):e70642. doi: 10.1111/tpj.v124.5 URL