RECENT ADVANCES IN THE REGULATION OF CLIMACTERIC FRUIT RIPENING: HORMONE, TRANSCRIPTION FACTOR AND EPIGENETIC MODIFICATIONS

Highlight

• The dynamic interplay between phytohormones plays an important part in climacteric fruit ripening.

• Transcription factors are critical for the regulation of climacteric fruit ripening.

• Epigenetic modifications act as important regulators of fruit ripening.

Abstract

Fruit ripening is a complex developmental process made up of genetically programmed physiological and biochemical activities. It culminates in desirable changes in the structural and textural properties and is governed by a complex regulatory network. Much is known about ethylene, one of the most important metabolites promoting the ripening of climacteric fruits. However, the dynamic interplay between phytohormones also plays an important part. Additional regulatory factors such as transcription factors (TFs) and epigenetic modifications also play vital role in the regulation of climacteric fruit ripening. Here, we review and evaluate the complex regulatory network comprising interactions between hormones and the action of TFs and epigenetic modifications during climacteric fruit ripening.

Keywords

climacteric fruit ripening / phytohormones / TFs / epigenetic modifications

参考文献

[1] Karlova R, Chapman N, David K, Angenent G C, Seymour G B, de Maagd R A. Transcriptional control of fleshy fruit development and ripening. Journal of Experimental Botany, 2014, 65(16): 4527–4541

[2] Giovannoni J, Nguyen C, Ampofo B, Zhong S, Fei Z. The epigenome and transcriptional dynamics of fruit ripening. Annual Review of Plant Biology, 2017, 68(1): 61–84

[3] Giovannoni J. Molecular biology of fruit maturation and ripening. Annual Review of Plant Physiology and Plant Molecular Biology, 2001, 52(1): 725–749

[4] Kevany B M, Tieman D M, Taylor M G, Cin V D, Klee H J. Ethylene receptor degradation controls the timing of ripening in tomato fruit. Plant Journal, 2007, 51(3): 458–467

[5] Barry C S, Giovannoni J J. Ripening in the tomato Green-ripe mutant is inhibited by ectopic expression of a protein that disrupts ethylene signaling. Proceedings of the National Academy of Sciences of the United States of America, 2006, 103(20): 7923–7928

[6] Manjunatha G, Gupta K J, Lokesh V, Mur L A, Neelwarne B. Nitric oxide counters ethylene effects on ripening fruits. Plant Signaling & Behavior, 2012, 7(4): 476–483

[7] Cherian S, Figueroa C R, Nair H. ‘Movers and shakers’ in the regulation of fruit ripening: a cross-dissection of climacteric versus non-climacteric fruit. Journal of Experimental Botany, 2014, 65(17): 4705–4722

[8] Giovannoni J J. Genetic regulation of fruit development and ripening. Plant Cell, 2004, 16(Suppl 1): S170–S180

[9] Wang W, Cai J, Wang P, Tian S, Qin G. Post-transcriptional regulation of fruit ripening and disease resistance in tomato by the vacuolar protease SlVPE3. Genome Biology, 2017, 18(1): 47

[10] Alexander L, Grierson D. Ethylene biosynthesis and action in tomato: a model for climacteric fruit ripening. Journal of Experimental Botany, 2002, 53(377): 2039–2055

[11] Fuentes L, Figueroa C R, Valdenegro M. Recent advances in hormonal regulation and cross-talk during non-climacteric fruit development and ripening. Horticulturae, 2019, 5(2): 45

[12] Li C, Jia H, Chai Y, Shen Y. Abscisic acid perception and signaling transduction in strawberry: a model for non-climacteric fruit ripening. Plant Signaling & Behavior, 2011, 6(12): 1950–1953

[13] Castellarin S D, Gambetta G A, Wada H, Shackel K A, Matthews M A. Fruit ripening in Vitis vinifera: spatiotemporal relationships among turgor, sugar accumulation, and anthocyanin biosynthesis. Journal of Experimental Botany, 2011, 62(12): 4345–4354

[14] Liu K, Yuan C, Feng S, Zhong S, Li H, Zhong J, Shen C, Liu J. Genome-wide analysis and characterization of Aux/IAA family genes related to fruit ripening in papaya (Carica papaya L.). BMC Genomics, 2017, 18(1): 351

[15] Pérez-Llorca M, Muñoz P, Müller M, Munné-Bosch S. Biosynthesis, metabolism and function of auxin, salicylic acid and melatonin in climacteric and non-climacteric fruits. Frontiers of Plant Science, 2019, 10: 136

[16] Chen Y, Grimplet J, David K, Castellarin S D, Terol J, Wong D C J, Luo Z, Schaffer R, Celton J M, Talon M, Gambetta G A, Chervin C. Ethylene receptors and related proteins in climacteric and non-climacteric fruits. Plant Science, 2018, 276: 63–72

[17] Symons G M, Chua Y J, Ross J J, Quittenden L J, Davies N W, Reid J B. Hormonal changes during non-climacteric ripening in strawberry. Journal of Experimental Botany, 2012, 63(13): 4741–4750

[18] Li Y, Lu Y, Li L, Chu Z, Zhang H, Li H, Fernie A R, Ouyang B. Impairment of hormone pathways results in a general disturbance of fruit primary metabolism in tomato. Food Chemistry, 2019, 274: 170–179

[19] Cheng J, Niu Q, Zhang B, Chen K, Yang R, Zhu J K, Zhang Y, Lang Z. Downregulation of RdDM during strawberry fruit ripening. Genome Biology, 2018, 19(1): 212

[20] Manning K, Tör M, Poole M, Hong Y, Thompson A J, King G J, Giovannoni J J, Seymour G B. A naturally occurring epigenetic mutation in a gene encoding an SBP-box transcription factor inhibits tomato fruit ripening. Nature Genetics, 2006, 38(8): 948–952

[21] Zeng Y, Pan Z, Wang L, Ding Y, Xu Q, Xiao S, Deng X. Phosphoproteomic analysis of chromoplasts from sweet orange during fruit ripening. Physiologia Plantarum, 2014, 150(2): 252–270

[22] Kamiyoshihara Y, Tieman D M, Huber D J, Klee H J. Ligand-induced alterations in the phosphorylation state of ethylene receptors in tomato fruit. Plant Physiology, 2012, 160(1): 488–497

[23] Guo J E, Hu Z, Yu X, Li A, Li F, Wang Y, Tian S, Chen G. A histone deacetylase gene, SlHDA3, acts as a negative regulator of fruit ripening and carotenoid accumulation. Plant Cell Reports, 2018, 37(1): 125–135

[24] Wang Y, Wang W, Cai J, Zhang Y, Qin G, Tian S. Tomato nuclear proteome reveals the involvement of specific E2 ubiquitin-conjugating enzymes in fruit ripening. Genome Biology, 2014, 15(12): 548

[25] Li S, Chen K, Grierson D. A critical evaluation of the role of ethylene and MADS transcription factors in the network controlling fleshy fruit ripening. New Phytologist, 2019, 221(4): 1724–1741

[26] Buesa C, Dominguez M, Vendrell M. Abscisic acid effects on ethylene production and respiration rate in detached apple fruits at different stages of development. Spanish Journal of Food Science and Technology, 1994, 34(5): 495–506

[27] Li T, Jiang Z, Zhang L, Tan D, Wei Y, Yuan H, Li T, Wang A. Apple (Malus domestica) MdERF2 negatively affects ethylene biosynthesis during fruit ripening by suppressing MdACS1 transcription. Plant Journal, 2016, 88(5): 735–748

[28] Li T, Xu Y, Zhang L, Ji Y, Tan D, Yuan H, Wang A. The jasmonate-activated transcription factor MdMYC2 regulates ETHYLENE RESPONSE FACTOR and ethylene biosynthetic genes to promote ethylene biosynthesis during apple fruit ripening. Plant Cell, 2017, 29(6): 1316–1334

[29] Jiang Y, Joyce D C, Macnish A J. Effect of abscisic acid on banana fruit ripening in relation to the role of ethylene. Journal of Plant Growth Regulation, 2000, 19(1): 106–111

[30] Guo Y F, Shan W, Liang S M, Wu C J, Wei W, Chen J Y, Lu W J, Kuang J F. MaBZR1/2 act as transcriptional repressors of ethylene biosynthetic genes in banana fruit. Physiologia Plantarum, 2019, 165(3): 555–568

[31] Khader S E S A, Singh B P, Khan S A. Effect of GA3 as a post-harvest treatment of mango fruit on ripening, amylase and peroxidase activity and quality during storage. Scientia Horticulturae, 1988, 36(3-4): 261–266

[32] Zaharah S S, Singh Z, Symons G M, Reid J B. Role of brassinosteroids, ethylene, abscisic acid, and indole-3-acetic acid in mango fruit ripening. Journal of Plant Growth Regulation, 2012, 31(3): 363–372

[33] Trainotti L, Tadiello A, Casadoro G. The involvement of auxin in the ripening of climacteric fruits comes of age: the hormone plays a role of its own and has an intense interplay with ethylene in ripening peaches. Journal of Experimental Botany, 2007, 58(12): 3299–3308

[34] Tatsuki M, Nakajima N, Fujii H, Shimada T, Nakano M, Hayashi K, Hayama H, Yoshioka H, Nakamura Y. Increased levels of IAA are required for system 2 ethylene synthesis causing fruit softening in peach (Prunus persica L. Batsch). Journal of Experimental Botany, 2013, 64(4): 1049–1059

[35] Shi H Y, Zhang Y X. Expression and regulation of pear 1-aminocyclopropane-1-carboxylic acid synthase gene (PpACS1a) during fruit ripening, under salicylic acid and indole-3-acetic acid treatment, and in diseased fruit. Molecular Biology Reports, 2014, 41(6): 4147–4154

[36] Yue P T, Wang Y N, Bu H D, Li X Y, Yuan H, Wang A D. Ethylene promotes IAA reduction through PuERFs-activated PuGH3.1 during fruit ripening in pear (Pyrus ussuriensis). Postharvest Biology and Technology, 2019, 157: 110955

[37] Khan A, Singh Z. Methyl jasmonate promotes fruit ripening and improves fruit quality in Japanese plum. Journal of Horticultural Science & Biotechnology, 2007, 82(5): 695–706

[38] Kende H. Ethylene biosynthesis. Annual Review of Plant Biology, 1993, 44(1): 283–307

[39] Payasi A, Sanwal G. Ripening of climacteric fruits and their control. Journal of Food Biochemistry, 2010, 34(4): 679–710

[40] Yue P, Lu Q, Liu Z, Lv T, Li X, Bu H, Liu W, Xu Y, Yuan H, Wang A. Auxin-activated MdARF5 induces the expression of ethylene biosynthetic genes to initiate apple fruit ripening. New Phytologist, 2020, 226(6): 1781–1795

[41] Liu K, Kang B C, Jiang H, Moore S L, Li H, Watkins C B, Setter T L, Jahn M M A. A GH3-like gene, CcGH3, isolated from Capsicum chinense L. fruit is regulated by auxin and ethylene. Plant Molecular Biology, 2005, 58(4): 447–464

[42] Zhang M, Leng P, Zhang G, Li X. Cloning and functional analysis of 9-cis-epoxycarotenoid dioxygenase (NCED) genes encoding a key enzyme during abscisic acid biosynthesis from peach and grape fruits. Journal of Plant Physiology, 2009, 166(12): 1241–1252

[43] Chen S, Wang X, Zhang L, Lin S, Liu D, Wang Q, Cai S, El-Tanbouly R, Gan L, Wu H, Li Y. Identification and characterization of tomato gibberellin 2-oxidases (GA2oxs) and effects of fruit-specific SlGA2ox1 overexpression on fruit and seed growth and development. Horticulture Research, 2016, 3(1): 16059

[44] Ben-Arie R, Saks Y, Sonego L, Frank A. Cell wall metabolism in gibberellin-treated persimmon fruits. Plant Growth Regulation, 1996, 19(1): 25–33

[45] Clouse S D. Brassinosteroid signal transduction: from receptor kinase activation to transcriptional networks regulating plant development. Plant Cell, 2011, 23(4): 1219–1230

[46] Saniewski M, Czapski J, Nowacki J, Lange E. The effect of methyl jasmonate on ethylene and l-aminocyclopropane-1-carboxylic acid production in apple fruits. Biologia Plantarum, 1987, 29(3): 199–203

[47] Davey J E, Van Staden J. Endogenous cytokinins in the fruits of ripening and non-ripening tomatoes. Plant Science Letters, 1978, 11(3-4): 359–364

[48] Barry C S, Llop-Tous M I, Grierson D. The regulation of 1-aminocyclopropane-1-carboxylic acid synthase gene expression during the transition from system-1 to system-2 ethylene synthesis in tomato. Plant Physiology, 2000, 123(3): 979–986

[49] Nakatsuka A, Murachi S, Okunishi H, Shiomi S, Nakano R, Kubo Y, Inaba A. Differential expression and internal feedback regulation of 1-aminocyclopropane-1-carboxylate synthase, 1-aminocyclopropane-1-carboxylate oxidase, and ethylene receptor genes in tomato fruit during development and ripening. Plant Physiology, 1998, 118(4): 1295–1305

[50] McMurchie E J, McGlasson W B, Eaks I L. Treatment of fruit with propylene gives information about the biogenesis of ethylene. Nature, 1972, 237(5352): 235–236

[51] Yang S F, Hoffman N E. Ethylene biosynthesis and its regulation in higher plants. Annual Review of Plant Physiology, 1984, 35(1): 155–189

[52] Seymour G B, Taylor J E, Tucker G A. Biochemistry of fruit ripening. Springer, 1993

[53] Lin Z, Zhong S, Grierson D. Recent advances in ethylene research. Journal of Experimental Botany, 2009, 60(12): 3311–3336

[54] Li T, Tan D, Liu Z, Jiang Z, Wei Y, Zhang L, Li X, Yuan H, Wang A. Apple MdACS6 regulates ethylene biosynthesis during fruit development involving ethylene-responsive factor. Plant & Cell Physiology, 2015, 56(10): 1909–1917

[55] Wang A, Yamakake J, Kudo H, Wakasa Y, Hatsuyama Y, Igarashi M, Kasai A, Li T, Harada T. Null mutation of the MdACS3 gene, coding for a ripening-specific 1-aminocyclopropane-1-carboxylate synthase, leads to long shelf life in apple fruit. Plant Physiology, 2009, 151(1): 391–399

[56] Varanasi V, Shin S, Mattheis J, Rudell D, Zhu Y M. Expression profiles of the MdACS3 gene suggest a function as an accelerator of apple (Malus × domestica) fruit ripening. Postharvest Biology and Technology, 2011, 62(2): 141–148

[57] Tan D M, Li T Z, Wang A D. Apple 1-aminocyclopropane-1-carboxylic acid synthase genes, MdACS1 and MdACS3a, are expressed in different systems of ethylene biosynthesis. Plant Molecular Biology Reporter, 2013, 31(1): 204–209

[58] Sunako T, Sakuraba W, Senda M, Akada S, Ishikawa R, Niizeki M, Harada T. An allele of the ripening-specific 1-aminocyclopropane-1-carboxylic acid synthase gene (ACS1) in apple fruit with a long storage life. Plant Physiology, 1999, 119(4): 1297–1304

[59] Dandekar A M, Teo G, Defilippi B G, Uratsu S L, Passey A J, Kader A A, Stow J R, Colgan R J, James D J. Effect of down-regulation of ethylene biosynthesis on fruit flavor complex in apple fruit. Transgenic Research, 2004, 13(4): 373–384

[60] Oeller P W, Lu M W, Taylor L P, Pike D A, Theologis A. Reversible inhibition of tomato fruit senescence by antisense RNA. Science, 1991, 254(5030): 437–439

[61] Barry C S, Blume B, Bouzayen M, Cooper W, Hamilton A J, Grierson D. Differential expression of the 1-aminocyclopropane-1-carboxylate oxidase gene family of tomato. Plant Journal, 1996, 9(4): 525–535

[62] Schaffer R J, Friel E N, Souleyre E J, Bolitho K, Thodey K, Ledger S, Bowen J H, Ma J H, Nain B, Cohen D, Gleave A P, Crowhurst R N, Janssen B J, Yao J L, Newcomb R D. A genomics approach reveals that aroma production in apple is controlled by ethylene predominantly at the final step in each biosynthetic pathway. Plant Physiology, 2007, 144(4): 1899–1912

[63] Jiang Y M, Fu J R. Ethylene regulation of fruit ripening: molecular aspects. Plant Growth Regulation, 2000, 30(3): 193–200

[64] Liu Y, Hoffman N E, Yang S F. Promotion by ethylene of the capability to convert 1-aminocyclopropane-1-carboxylic acid to ethylene in preclimacteric tomato and cantaloupe fruits. Plant Physiology, 1985, 77(2): 407–411

[65] Guo H, Ecker J R. The ethylene signaling pathway: new insights. Current Opinion in Plant Biology, 2004, 7(1): 40–49

[66] Ireland H S, Guillen F, Bowen J H, Tacken E J, Putterill J, Schaffer R J, Johnston J W. Mining the apple genome reveals a family of nine ethylene receptor genes. Postharvest Biology and Technology, 2012, 72: 42–46

[67] Wilkinson J Q, Lanahan M B, Yen H C, Giovannoni J J, Klee H J. An ethylene-inducible component of signal transduction encoded by never-ripe. Science, 1995, 270(5243): 1807–1809

[68] Zhou D, Kalaitzís P, Mattoo A K, Tucker M L. The mRNA for an ETR1 homologue in tomato is constitutively expressed in vegetative and reproductive tissues. Plant Molecular Biology, 1996, 30(6): 1331–1338

[69] Lashbrook C C, Tieman D M, Klee H J. Differential regulation of the tomato ETR gene family throughout plant development. Plant Journal, 1998, 15(2): 243–252

[70] Tieman D M, Klee H J. Differential expression of two novel members of the tomato ethylene-receptor family. Plant Physiology, 1999, 120(1): 165–172

[71] Gao Z, Chen Y F, Randlett M D, Zhao X C, Findell J L, Kieber J J, Schaller G E. Localization of the Raf-like kinase CTR1 to the endoplasmic reticulum of Arabidopsis through participation in ethylene receptor signaling complexes. Journal of Biological Chemistry, 2003, 278(36): 34725–34732

[72] Huang Y, Li H, Hutchison C E, Laskey J, Kieber J J. Biochemical and functional analysis of CTR1, a protein kinase that negatively regulates ethylene signaling in Arabidopsis. Plant Journal, 2003, 33(2): 221–233

[73] Leclercq J, Adams-Phillips L C, Zegzouti H, Jones B, Latché A, Giovannoni J J, Pech J C, Bouzayen M. LeCTR1, a tomato CTR1-like gene, demonstrates ethylene signaling ability in Arabidopsis and novel expression patterns in tomato. Plant Physiology, 2002, 130(3): 1132–1142

[74] Adams-Phillips L, Barry C, Kannan P, Leclercq J, Bouzayen M, Giovannoni J. Evidence that CTR1-mediated ethylene signal transduction in tomato is encoded by a multigene family whose members display distinct regulatory features. Plant Molecular Biology, 2004, 54(3): 387–404

[75] Alonso J M, Stepanova A N. The ethylene signaling pathway. Science, 2004, 306(5701): 1513–1515

[76] Yin X R, Allan A C, Chen K S, Ferguson I B. Kiwifruit EIL and ERF genes involved in regulating fruit ripening. Plant Physiology, 2010, 153(3): 1280–1292

[77] Nakano T, Suzuki K, Fujimura T, Shinshi H. Genome-wide analysis of the ERF gene family in Arabidopsis and rice. Plant Physiology, 2006, 140(2): 411–432

[78] Ohta M, Matsui K, Hiratsu K, Shinshi H, Ohme-Takagi M. Repression domains of class II ERF transcriptional repressors share an essential motif for active repression. Plant Cell, 2001, 13(8): 1959–1968

[79] Ohta M, Ohme-Takagi M, Shinshi H. Three ethylene-responsive transcription factors in tobacco with distinct transactivation functions. Plant Journal, 2000, 22(1): 29–38

[80] Fujimoto S Y, Ohta M, Usui A, Shinshi H, Ohme-Takagi M. Arabidopsis ethylene-responsive element binding factors act as transcriptional activators or repressors of GCC box-mediated gene expression. Plant Cell, 2000, 12(3): 393–404

[81] Zhang Z, Zhang H, Quan R, Wang X C, Huang R. Transcriptional regulation of the ethylene response factor LeERF2 in the expression of ethylene biosynthesis genes controls ethylene production in tomato and tobacco. Plant Physiology, 2009, 150(1): 365–377

[82] Xiao Y Y, Chen J Y, Kuang J F, Shan W, Xie H, Jiang Y M, Lu W J. Banana ethylene response factors are involved in fruit ripening through their interactions with ethylene biosynthesis genes. Journal of Experimental Botany, 2013, 64(8): 2499–2510

[83] Kumar R, Khurana A, Sharma A K. Role of plant hormones and their interplay in development and ripening of fleshy fruits. Journal of Experimental Botany, 2014, 65(16): 4561–4575

[84] Abel S, Theologis A. Early genes and auxin action. Plant Physiology, 1996, 111(1): 9–17

[85] Buta J G, Spaulding D W. Changes in indole-3-acetic acid and abscisic acid levels during tomato (Lycopersicon esculentum Mill.) fruit development and ripening. Journal of Plant Growth Regulation, 1994, 13(3): 163–166

[86] Mapelli S, Frova C, Torti G, Soressi G P. Relationship between set, development and activities of growth regulators in tomato fruits. Plant & Cell Physiology, 1978, 19(7): 1281–1288

[87] Kumar R, Agarwal P, Tyagi A K, Sharma A K. Genome-wide investigation and expression analysis suggest diverse roles of auxin-responsive GH3 genes during development and response to different stimuli in tomato (Solanum lycopersicum). Molecular Genetics and Genomics, 2012, 287(3): 221–235

[88] Miller A N, Walsh C S, Cohen J D. Measurement of indole-3-acetic acid in peach fruits (Prunus persica L. Batsch cv Redhaven) during development. Plant Physiology, 1987, 84(2): 491–494

[89] Lavy M, Estelle M. Mechanisms of auxin signaling. Development, 2016, 143(18): 3226–3229

[90] Leyser O. Auxin signaling. Plant Physiology, 2018, 176(1): 465–479

[91] Lohani S, Trivedi P K, Nath P. Changes in activities of cell wall hydrolases during ethylene-induced ripening in banana: effect of 1-MCP, ABA and IAA. Postharvest Biology and Technology, 2004, 31(2): 119–126

[92] Li J, Tao X, Li L, Mao L, Luo Z, Khan Z U, Ying T. Comprehensive RNA-Seq analysis on the regulation of tomato ripening by exogenous auxin. PLoS One, 2016, 11(5): e0156453

[93] El-Sharkawy I, Sherif S M, Jones B, Mila I, Kumar P P, Bouzayen M, Jayasankar S. TIR1-like auxin-receptors are involved in the regulation of plum fruit development. Journal of Experimental Botany, 2014, 65(18): 5205–5215

[94] Bleecker A B, Kende H. Ethylene: a gaseous signal molecule in plants. Annual Review of Cell and Developmental Biology, 2000, 16(1): 1–18

[95] Desai B B, Deshpande P B. Chemical control of ripening in banana. Physiologia Plantarum, 1978, 44(3): 238–240

[96] McAtee P, Karim S, Schaffer R, David K. A dynamic interplay between phytohormones is required for fruit development, maturation, and ripening. Frontiers of Plant Science, 2013, 4: 79

[97] Jia H F, Chai Y M, Li C L, Lu D, Luo J J, Qin L, Shen Y Y. Abscisic acid plays an important role in the regulation of strawberry fruit ripening. Plant Physiology, 2011, 157(1): 188–199

[98] Hou B Z, Li C L, Han Y Y, Shen Y Y. Characterization of the hot pepper (Capsicum frutescens) fruit ripening regulated by ethylene and ABA. BMC Plant Biology, 2018, 18(1): 162

[99] Mou W, Li D, Luo Z, Li L, Mao L, Ying T. SlAREB1 transcriptional activation of NOR is involved in abscisic acid-modulated ethylene biosynthesis during tomato fruit ripening. Plant Science, 2018, 276: 239–249

[100] Wang Y, Wang Y, Ji K, Dai S, Hu Y, Sun L, Li Q, Chen P, Sun Y, Duan C, Wu Y, Luo H, Zhang D, Guo Y, Leng P. The role of abscisic acid in regulating cucumber fruit development and ripening and its transcriptional regulation. Plant Physiology and Biochemistry, 2013, 64: 70–79

[101] Zhang M, Yuan B, Leng P. The role of ABA in triggering ethylene biosynthesis and ripening of tomato fruit. Journal of Experimental Botany, 2009, 60(6): 1579–1588

[102] Zaharah S S, Singh Z, Symons G M, Reid J B. Mode of action of abscisic acid in triggering ethylene biosynthesis and softening during ripening in mango fruit. Postharvest Biology and Technology, 2013, 75: 37–44

[103] Vendrell M, Buesa C. Relationship between abscisic acid content and ripening of apples. In: Herregods M, ed. International symposium on postharvest handling of fruit and vegetables. Acta Horticulturae, 1989, (258): 45

[104] Shu K, Liu X D, Xie Q, He Z H. Two faces of one seed: hormonal regulation of dormancy and germination. Molecular Plant, 2016, 9(1): 34–45

[105] Pharis R P, King R W. Gibberellins and reproductive development in seed plants. Annual Review of Plant Physiology, 1985, 36(1): 517–568

[106] Serrani J C, Sanjuán R, Ruiz-Rivero O, Fos M, García-Martínez J L. Gibberellin regulation of fruit set and growth in tomato. Plant Physiology, 2007, 145(1): 246–257

[107] Li H, Wu H, Qi Q, Li H, Li Z, Chen S, Ding Q, Wang Q, Yan Z, Gai Y, Jiang X, Ding J, Gu T, Hou X, Richard M, Zhao Y, Li Y. Gibberellins play a role in regulating tomato fruit ripening. Plant & Cell Physiology, 2019, 60(7): 1619–1629

[108] Srivastava A, Handa A K. Hormonal regulation of tomato fruit development: a molecular perspective. Journal of Plant Growth Regulation, 2005, 24(2): 67–82

[109] Dostal H C, Leopold A C. Gibberellin delays ripening of tomatoes. Science, 1967, 158(3808): 1579–1580

[110] Mandava N B. Plant growth-promoting brassinosteroids. Annual Review of Plant Physiology and Plant Molecular Biology, 1988, 39(1): 23–52

[111] Montoya T, Nomura T, Yokota T, Farrar K, Harrison K, Jones J D G, Kaneta T, Kamiya Y, Szekeres M, Bishop G J. Patterns of Dwarf expression and brassinosteroid accumulation in tomato reveal the importance of brassinosteroid synthesis during fruit development. Plant Journal, 2005, 42(2): 262–269

[112] Vidya Vardhini B, Rao S S R. Acceleration of ripening of tomato pericarp discs by brassinosteroids. Phytochemistry, 2002, 61(7): 843–847

[113] Zhu T, Tan W R, Deng X G, Zheng T, Zhang D W, Lin H H. Effects of brassinosteroids on quality attributes and ethylene synthesis in postharvest tomato fruit. Postharvest Biology and Technology, 2015, 100: 196–204

[114] Ji Y L Y, Qu Z Y, Jiang J J, Yan J F, Chu M Y, Xu X, Su H, Yuan A D, Wang. The mechanism for brassinosteroids suppressing climacteric fruit ripening. Plant Physiology, 2021: kiab013 doi:10.1093/plphys/kiab013

[115] Li X J, Chen X J, Guo X, Yin L L, Ahammed G J, Xu C J, Chen K S, Liu C C, Xia X J, Shi K, Zhou J, Zhou Y H, Yu J Q. DWARF overexpression induces alteration in phytohormone homeostasis, development, architecture and carotenoid accumulation in tomato. Plant Biotechnology Journal, 2016, 14(3): 1021–1033

[116] Nie S, Huang S, Wang S, Cheng D, Liu J, Lv S, Li Q, Wang X. Enhancing brassinosteroid signaling via overexpression of tomato (Solanum lycopersicum) SlBRI1 improves major agronomic traits. Frontiers of Plant Science, 2017, 8: 1386

[117] Zhu Z, Zhang Z Q, Qin G Z, Tian S P. Effects of brassinosteroids on postharvest disease and senescence of jujube fruit in storage. Postharvest Biology and Technology, 2010, 56(1): 50–55

[118] Lv B, Tian H, Zhang F, Liu J, Lu S, Bai M, Li C, Ding Z. Brassinosteroids regulate root growth by controlling reactive oxygen species homeostasis and dual effect on ethylene synthesis in Arabidopsis. PLOS Genetics, 2018, 14(1): e1007144

[119] Wasternack C, Song S. Jasmonates: biosynthesis, metabolism, and signaling by proteins activating and repressing transcription. Journal of Experimental Botany, 2017, 68(6): 1303–1321

[120] Wasternack C, Hause B. Jasmonates: biosynthesis, perception, signal transduction and action in plant stress response, growth and development. An update to the 2007 review in Annals of Botany. Annals of Botany, 2013, 111(6): 1021–1058

[121] Barry C S, Giovannoni J J. Ethylene and fruit ripening. Journal of Plant Growth Regulation, 2007, 26(2): 143–159

[122] Kondo S, Tomiyama A, Seto H. Changes of endogenous jasmonic acid and methyl jasmonate in apples and sweet cherries during fruit development. Journal of the American Society for Horticultural Science, 2000, 125(3): 282–287

[123] Saniewski M, Miszczak A, Kawa-Miszczak L, Wegrzynowicz-Lesiak E, Miyamoto K, Ueda J. Effects of methyl jasmonate on anthocyanin accumulation, ethylene production, and CO₂ evolution in uncooled and cooled tulip bulbs. Journal of Plant Growth Regulation, 1998, 17(1): 33–37

[124] Lalel H J D, Singh Z, Tan S C. The role of methyl jasmonate in mango ripening and biosynthesis of aroma volatile compounds. Journal of Horticultural Science & Biotechnology, 2003, 78(4): 470–484

[125] Saniewski M, Czapski J, Nowacki J. Relationship between stimulatory effect of methyl jasmonate on ethylene production and 1-aminocyclopropane-1-carboxylic acid content in tomatoes. Biologia Plantarum, 1987, 29(1): 17–21

[126] Kazan K, Manners J M. MYC2: the master in action. Molecular Plant, 2013, 6(3): 686–703

[127] Fernández-Calvo P, Chini A, Fernández-Barbero G, Chico J M, Gimenez-Ibanez S, Geerinck J, Eeckhout D, Schweizer F, Godoy M, Franco-Zorrilla J M, Pauwels L, Witters E, Puga M I, Paz-Ares J, Goossens A, Reymond P, De Jaeger G, Solano R. The Arabidopsis bHLH transcription factors MYC3 and MYC4 are targets of JAZ repressors and act additively with MYC2 in the activation of jasmonate responses. Plant Cell, 2011, 23(2): 701–715

[128] Zhang X, Zhu Z, An F, Hao D, Li P, Song J, Yi C, Guo H. Jasmonate-activated MYC2 represses ETHYLENE INSENSITIVE3 activity to antagonize ethylene-promoted apical hook formation in Arabidopsis. Plant Cell, 2014, 26(3): 1105–1117

[129] Rudell D R, Fellman J K, Mattheis J P. Preharvest application of methyl jasmonate to ‘Fuji’ apples enhances red coloration and affects fruit size, splitting, and bitter pit incidence. HortScience, 2005, 40(6): 1760–1762

[130] Liu W, Li T, Yuan H, Tan D, Wang A. Enhancement of apple coloration using jasmonate treatment without sacrificing storage potential. Plant Signaling & Behavior, 2018, 13(1): e1422467

[131] Kieber J J, Schaller G E. Cytokinins. In: The Arabidopsis Book. The American Society of Plant Biologists, 2014, 12

[132] Varga A, Bruinsma J J. The growth and ripening of tomato fruits at different levels of endogenous cytokinins. Journal of Horticultural Science, 1974, 49(2): 135–142

[133] Desai N, Chism G M. Changes in cytokinin activity in the ripening tomato fruit. Journal of Food Science, 1978, 43(4): 1324–1326

[134] Ainalidou A, Tanou G, Belghazi M, Samiotaki M, Diamantidis G, Molassiotis A, Karamanoli K. Integrated analysis of metabolites and proteins reveal aspects of the tissue-specific function of synthetic cytokinin in kiwifruit development and ripening. Journal of Proteomics, 2016, 143: 318–333

[135] Setha S. Roles of abscisic acid in fruit ripening. Walailak Journal of Science and Technology, 2012, 9(4): 297–308

[136] Sun L, Sun Y, Zhang M, Wang L, Ren J, Cui M, Wang Y, Ji K, Li P, Li Q, Chen P, Dai S, Duan C, Wu Y, Leng P. Suppression of 9-cis-epoxycarotenoid dioxygenase, which encodes a key enzyme in abscisic acid biosynthesis, alters fruit texture in transgenic tomato. Plant Physiology, 2012, 158(1): 283–298

[137] Jones B, Frasse P, Olmos E, Zegzouti H, Li Z G, Latché A, Pech J C, Bouzayen M. Down-regulation of DR12, an auxin-response-factor homolog, in the tomato results in a pleiotropic phenotype including dark green and blotchy ripening fruit. Plant Journal, 2002, 32(4): 603–613

[138] Parra-Lobato M C, Gomez-Jimenez M C. Polyamine-induced modulation of genes involved in ethylene biosynthesis and signalling pathways and nitric oxide production during olive mature fruit abscission. Journal of Experimental Botany, 2011, 62(13): 4447–4465

[139] Torrigiani P, Bressanin D, Beatriz Ruiz K, Tadiello A, Trainotti L, Bonghi C, Ziosi V, Costa G. Spermidine application to young developing peach fruits leads to a slowing down of ripening by impairing ripening-related ethylene and auxin metabolism and signaling. Physiologia Plantarum, 2012, 146(1): 86–98

[140] Li N, Parsons B L, Liu D R, Mattoo A K. Accumulation of wound-inducible ACC synthase transcript in tomato fruit is inhibited by salicylic acid and polyamines. Plant Molecular Biology, 1992, 18(3): 477–487

[141] Rümer S, Gupta K J, Kaiser W M. Plant cells oxidize hydroxylamines to NO. Journal of Experimental Botany, 2009, 60(7): 2065–2072

[142] Manjunatha G, Lokesh V, Neelwarne B. Nitric oxide in fruit ripening: trends and opportunities. Biotechnology Advances, 2010, 28(4): 489–499

[143] Ziosi V, Bonghi C, Bregoli A M, Trainotti L, Biondi S, Sutthiwal S, Kondo S, Costa G, Torrigiani P. Jasmonate-induced transcriptional changes suggest a negative interference with the ripening syndrome in peach fruit. Journal of Experimental Botany, 2008, 59(3): 563–573

[144] Fan X T, Mattheis J P, Fellman J K. A role for jasmonates in climacteric fruit ripening. Planta, 1998, 204(4): 444–449

[145] Singh R, Singh P, Pathak N, Singh V K, Dwivedi U N. Modulation of mango ripening by chemicals: physiological and biochemical aspects. Plant Growth Regulation, 2007, 53(2): 137–145

[146] Martínez Romero D, Valero D, Serrano M, Burló F, Carbonell A, Burgos L, Riquelme F. Exogenous polyamines and gibberellic acid effects on peach (Prunus persica L.) storability improvement. Journal of Food Science, 2000, 65(2): 288–294

[147] Vrebalov J, Ruezinsky D, Padmanabhan V, White R, Medrano D, Drake R, Schuch W, Giovannoni J. A MADS-box gene necessary for fruit ripening at the tomato ripening-inhibitor (rin) locus. Science, 2002, 296(5566): 343–346

[148] Ito Y, Nishizawa-Yokoi A, Endo M, Mikami M, Shima Y, Nakamura N, Kotake-Nara E, Kawasaki S, Toki S. Re-evaluation of the rin mutation and the role of RIN in the induction of tomato ripening. Nature Plants, 2017, 3(11): 866–874

[149] Li S, Xu H, Ju Z, Cao D, Zhu H, Fu D, Grierson D, Qin G, Luo Y, Zhu B. The RIN-MC fusion of MADS-box transcription factors has transcriptional activity and modulates expression of many ripening genes. Plant Physiology, 2018, 176(1): 891–909

[150] Li S, Zhu B, Pirrello J, Xu C, Zhang B, Bouzayen M, Chen K, Grierson D. Roles of RIN and ethylene in tomato fruit ripening and ripening-associated traits. New Phytologist, 2020, 226(2): 460–475

[151] Ito Y, Kitagawa M, Ihashi N, Yabe K, Kimbara J, Yasuda J, Ito H, Inakuma T, Hiroi S, Kasumi T. DNA-binding specificity, transcriptional activation potential, and the rin mutation effect for the tomato fruit-ripening regulator RIN. Plant Journal, 2008, 55(2): 212–223

[152] Fujisawa M, Nakano T, Ito Y. Identification of potential target genes for the tomato fruit-ripening regulator RIN by chromatin immunoprecipitation. BMC Plant Biology, 2011, 11(1): 26

[153] Martel C, Vrebalov J, Tafelmeyer P, Giovannoni J J. The tomato MADS-box transcription factor RIPENING INHIBITOR interacts with promoters involved in numerous ripening processes in a COLORLESS NONRIPENING-dependent manner. Plant Physiology, 2011, 157(3): 1568–1579

[154] Qin G, Wang Y, Cao B, Wang W, Tian S. Unraveling the regulatory network of the MADS box transcription factor RIN in fruit ripening. Plant Journal, 2012, 70(2): 243–255

[155] Gao C, Ju Z, Cao D, Zhai B, Qin G, Zhu H, Fu D, Luo Y, Zhu B. MicroRNA profiling analysis throughout tomato fruit development and ripening reveals potential regulatory role of RIN on microRNAs accumulation. Plant Biotechnology Journal, 2015, 13(3): 370–382

[156] Itkin M, Seybold H, Breitel D, Rogachev I, Meir S, Aharoni A. TOMATO AGAMOUS-LIKE 1 is a component of the fruit ripening regulatory network. Plant Journal, 2009, 60(6): 1081–1095

[157] Vrebalov J, Pan I L, Arroyo A J M, McQuinn R, Chung M, Poole M, Rose J, Seymour G, Grandillo S, Giovannoni J, Irish V F. Fleshy fruit expansion and ripening are regulated by the tomato SHATTERPROOF gene TAGL1. Plant Cell, 2009, 21(10): 3041–3062

[158] Bemer M, Karlova R, Ballester A R, Tikunov Y M, Bovy A G, Wolters-Arts M, Rossetto P B, Angenent G C, de Maagd R A. The tomato FRUITFULL homologs TDR4/FUL1 and MBP7/FUL2 regulate ethylene-independent aspects of fruit ripening. Plant Cell, 2012, 24(11): 4437–4451

[159] Fujisawa M, Shima Y, Nakagawa H, Kitagawa M, Kimbara J, Nakano T, Kasumi T, Ito Y. Transcriptional regulation of fruit ripening by tomato FRUITFULL homologs and associated MADS box proteins. Plant Cell, 2014, 26(1): 89–101

[160] Dong T, Hu Z, Deng L, Wang Y, Zhu M, Zhang J, Chen G. A tomato MADS-box transcription factor, SlMADS1, acts as a negative regulator of fruit ripening. Plant Physiology, 2013, 163(2): 1026–1036

[161] Xie Q, Hu Z, Zhu Z, Dong T, Zhao Z, Cui B, Chen G. Overexpression of a novel MADS-box gene SlFYFL delays senescence, fruit ripening and abscission in tomato. Scientific Reports, 2014, 4(1): 4367

[162] Giménez E, Pineda B, Capel J, Antón M T, Atarés A, Pérez-Martín F, García-Sogo B, Angosto T, Moreno V, Lozano R. Functional analysis of the Arlequin mutant corroborates the essential role of the Arlequin/TAGL1 gene during reproductive development of tomato. PLoS One, 2010, 5(12): e14427

[163] Shima Y, Kitagawa M, Fujisawa M, Nakano T, Kato H, Kimbara J, Kasumi T, Ito Y. Tomato FRUITFULL homologues act in fruit ripening via forming MADS-box transcription factor complexes with RIN. Plant Molecular Biology, 2013, 82(4-5): 427–438

[164] Hileman L C, Sundstrom J F, Litt A, Chen M, Shumba T, Irish V F. Molecular and phylogenetic analyses of the MADS-box gene family in tomato. Molecular Biology and Evolution, 2006, 23(11): 2245–2258

[165] Pirrello J, Prasad B C N, Zhang W, Chen K, Mila I, Zouine M, Latché A, Pech J C, Ohme-Takagi M, Regad F, Bouzayen M. Functional analysis and binding affinity of tomato ethylene response factors provide insight on the molecular bases of plant differential responses to ethylene. BMC Plant Biology, 2012, 12(1): 190

[166] Meng X, Xu J, He Y, Yang K Y, Mordorski B, Liu Y, Zhang S. Phosphorylation of an ERF transcription factor by Arabidopsis MPK3/MPK6 regulates plant defense gene induction and fungal resistance. Plant Cell, 2013, 25(3): 1126–1142

[167] Rodrigues M A, Bianchetti R E, Freschi L. Shedding light on ethylene metabolism in higher plants. Frontiers of Plant Science, 2014, 5: 665

[168] Chung M Y, Vrebalov J, Alba R, Lee J, McQuinn R, Chung J D, Klein P, Giovannoni J. A tomato (Solanum lycopersicum) APETALA2/ERF gene, SlAP2a, is a negative regulator of fruit ripening. Plant Journal, 2010, 64(6): 936–947

[169] Lee J M, Joung J G, McQuinn R, Chung M Y, Fei Z, Tieman D, Klee H, Giovannoni J. Combined transcriptome, genetic diversity and metabolite profiling in tomato fruit reveals that the ethylene response factor SlERF6 plays an important role in ripening and carotenoid accumulation. Plant Journal, 2012, 70(2): 191–204

[170] Karlova R, Rosin F M, Busscher-Lange J, Parapunova V, Do P T, Fernie A R, Fraser P D, Baxter C, Angenent G C, de Maagd R A. Transcriptome and metabolite profiling show that APETALA2a is a major regulator of tomato fruit ripening. Plant Cell, 2011, 23(3): 923–941

[171] Aida M, Ishida T, Fukaki H, Fujisawa H, Tasaka M. Genes involved in organ separation in Arabidopsis: an analysis of the cup-shaped cotyledon mutant. Plant Cell, 1997, 9(6): 841–857

[172] Olsen A N, Ernst H A, Leggio L L, Skriver K. NAC transcription factors: structurally distinct, functionally diverse. Trends in Plant Science, 2005, 10(2): 79–87

[173] Zhang H, Jin J, Tang L, Zhao Y, Gu X, Gao G, Luo J. PlantTFDB 2.0: update and improvement of the comprehensive plant transcription factor database. Nucleic Acids Research, 2011, 39(Database issue suppl_1): D1114–D1117

[174] Hao Y J, Wei W, Song Q X, Chen H W, Zhang Y Q, Wang F, Zou H F, Lei G, Tian A G, Zhang W K, Ma B, Zhang J S, Chen S Y. Soybean NAC transcription factors promote abiotic stress tolerance and lateral root formation in transgenic plants. Plant Journal, 2011, 68(2): 302–313

[175] Zhu M, Chen G, Zhou S, Tu Y, Wang Y, Dong T, Hu Z. A new tomato NAC (NAM/ATAF1/2/CUC2) transcription factor, SlNAC4, functions as a positive regulator of fruit ripening and carotenoid accumulation. Plant & Cell Physiology, 2014, 55(1): 119–135

[176] Yang S D, Seo P J, Yoon H K, Park C M. The Arabidopsis NAC transcription factor VNI2 integrates abscisic acid signals into leaf senescence via the COR/RD genes. Plant Cell, 2011, 23(6): 2155–2168

[177] Lindemose S, Jensen M K, de Velde J V, O’Shea C, Heyndrickx K S, Workman C T, Vandepoele K, Skriver K, Masi F D. A DNA-binding-site landscape and regulatory network analysis for NAC transcription factors in Arabidopsis thaliana. Nucleic Acids Research, 2014, 42(12): 7681–7693

[178] Moore S, Vrebalov J, Payton P, Giovannoni J. Use of genomics tools to isolate key ripening genes and analyse fruit maturation in tomato. Journal of Experimental Botany, 2002, 53(377): 2023–2030

[179] Osorio S, Alba R, Damasceno C M B, Lopez-Casado G, Lohse M, Zanor M I, Tohge T, Usadel B, Rose J K C, Fei Z, Giovannoni J J, Fernie A R. Systems biology of tomato fruit development: combined transcript, protein, and metabolite analysis of tomato transcription factor (nor, rin) and ethylene receptor (Nr) mutants reveals novel regulatory interactions. Plant Physiology, 2011, 157(1): 405–425

[180] Ma N, Feng H, Meng X, Li D, Yang D, Wu C, Meng Q. Overexpression of tomato SlNAC1 transcription factor alters fruit pigmentation and softening. BMC Plant Biology, 2014, 14(1): 351

[181] Meng C, Yang D, Ma X, Zhao W, Liang X, Ma N, Meng Q. Suppression of tomato SlNAC1 transcription factor delays fruit ripening. Journal of Plant Physiology, 2016, 193: 88–96

[182] Gao Y, Wei W, Zhao X, Tan X, Fan Z, Zhang Y, Jing Y, Meng L, Zhu B, Zhu H, Chen J, Jiang C Z, Grierson D, Luo Y, Fu D Q. A NAC transcription factor, NOR-like1, is a new positive regulator of tomato fruit ripening. Horticulture Research, 2018, 5(1): 75

[183] Lin Z, Hong Y, Yin M, Li C, Zhang K, Grierson D. A tomato HD-Zip homeobox protein, LeHB-1, plays an important role in floral organogenesis and ripening. Plant Journal, 2008, 55(2): 301–310

[184] Lovisetto A, Guzzo F, Tadiello A, Confortin E, Pavanello A, Botton A, Casadoro G. Characterization of a bZIP gene highly expressed during ripening of the peach fruit. Plant Physiology and Biochemistry, 2013, 70: 462–470

[185] Feller A, Machemer K, Braun E L, Grotewold E. Evolutionary and comparative analysis of MYB and bHLH plant transcription factors. Plant Journal, 2011, 66(1): 94–116

[186] Yao G, Ming M, Allan A C, Gu C, Li L, Wu X, Wang R, Chang Y, Qi K, Zhang S, Wu J. Map-based cloning of the pear gene MYB114 identifies an interaction with other transcription factors to coordinately regulate fruit anthocyanin biosynthesis. Plant Journal, 2017, 92(3): 437–451

[187] An X H, Tian Y, Chen K Q, Liu X J, Liu D D, Xie X B, Cheng C G, Cong P H, Hao Y J. MdMYB9 and MdMYB11 are involved in the regulation of the JA-induced biosynthesis of anthocyanin and proanthocyanidin in apples. Plant & Cell Physiology, 2015, 56(4): 650–662

[188] Fan Z Q, Ba L J, Shan W, Xiao Y Y, Lu W J, Kuang J F, Chen J Y. A banana R2R3-MYB transcription factor MaMYB3 is involved in fruit ripening through modulation of starch degradation by repressing starch degradation-related genes and MabHLH6. Plant Journal, 2018, 96(6): 1191–1205

[189] Kaufmann K, Pajoro A, Angenent G C. Regulation of transcription in plants: mechanisms controlling developmental switches. Nature Reviews: Genetics, 2010, 11(12): 830–842

[190] Feng S, Jacobsen S E, Reik W. Epigenetic reprogramming in plant and animal development. Science, 2010, 330(6004): 622–627

[191] Wollmann H, Berger F. Epigenetic reprogramming during plant reproduction and seed development. Current Opinion in Plant Biology, 2012, 15(1): 63–69

[192] Law J A, Jacobsen S E. Establishing, maintaining and modifying DNA methylation patterns in plants and animals. Nature Reviews: Genetics, 2010, 11(3): 204–220

[193] He X J, Chen T, Zhu J K. Regulation and function of DNA methylation in plants and animals. Cell Research, 2011, 21(3): 442–465

[194] Zhong S, Fei Z, Chen Y R, Zheng Y, Huang M, Vrebalov J, McQuinn R, Gapper N, Liu B, Xiang J, Shao Y, Giovannoni J J. Single-base resolution methylomes of tomato fruit development reveal epigenome modifications associated with ripening. Nature Biotechnology, 2013, 31(2): 154–159

[195] Teyssier E, Bernacchia G, Maury S, How Kit A, Stammitti-Bert L, Rolin D, Gallusci P. Tissue dependent variations of DNA methylation and endoreduplication levels during tomato fruit development and ripening. Planta, 2008, 228(3): 391–399

[196] Tate P H, Bird A P. Effects of DNA methylation on DNA-binding proteins and gene expression. Current Opinion in Genetics & Development, 1993, 3(2): 226–231

[197] Liu R, How-Kit A, Stammitti L, Teyssier E, Rolin D, Mortain-Bertrand A, Halle S, Liu M, Kong J, Wu C, Degraeve-Guibault C, Chapman N H, Maucourt M, Hodgman T C, Tost J, Bouzayen M, Hong Y, Seymour G B, Giovannoni J J, Gallusci P. A DEMETER-like DNA demethylase governs tomato fruit ripening. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(34): 10804–10809

[198] Lang Z, Wang Y, Tang K, Tang D, Datsenka T, Cheng J, Zhang Y, Handa A K, Zhu J K. Critical roles of DNA demethylation in the activation of ripening-induced genes and inhibition of ripening-repressed genes in tomato fruit. Proceedings of the National Academy of Sciences of the United States of America, 2017, 114(22): E4511–E4519

[199] Zhou L, Tian S, Qin G. RNA methylomes reveal the m⁶A-mediated regulation of DNA demethylase gene SlDML2 in tomato fruit ripening. Genome Biology, 2019, 20(1): 156

[200] Liu C, Lu F, Cui X, Cao X. Histone methylation in higher plants. Annual Review of Plant Biology, 2010, 61(1): 395–420

[201] Lü P, Yu S, Zhu N, Chen Y R, Zhou B, Pan Y, Tzeng D, Fabi J P, Argyris J, Garcia-Mas J, Ye N, Zhang J, Grierson D, Xiang J, Fei Z, Giovannoni J, Zhong S. Genome encode analyses reveal the basis of convergent evolution of fleshy fruit ripening. Nature Plants, 2018, 4(10): 784–791

[202] Liang Q, Deng H, Li Y, Liu Z, Shu P, Fu R, Zhang Y, Pirrello J, Zhang Y, Grierson D, Bouzayen M, Liu Y, Liu M. Like heterochromatin protein 1b represses fruit ripening via regulating the H3K27me3 levels in ripening-related genes in tomato. New Phytologist, 2020, 227(2): 485–497

[203] Li Z, Jiang G, Liu X, Ding X, Zhang D, Wang X, Zhou Y, Yan H, Li T, Wu K, Jiang Y, Duan X. Histone demethylase SlJMJ6 promotes fruit ripening by removing H3K27 methylation of ripening-related genes in tomato. New Phytologist, 2020, 227(4): 1138–1156

[204] Wang Z, Cao H, Chen F, Liu Y. The roles of histone acetylation in seed performance and plant development. Plant Physiology and Biochemistry, 2014, 84: 125–133

[205] Liu X, Yang S, Zhao M, Luo M, Yu C W, Chen C Y, Tai R, Wu K. Transcriptional repression by histone deacetylases in plants. Molecular Plant, 2014, 7(5): 764–772

[206] Aiese Cigliano R, Sanseverino W, Cremona G, Ercolano M R, Conicella C, Consiglio F M. Genome-wide analysis of histone modifiers in tomato: gaining an insight into their developmental roles. BMC Genomics, 2013, 14(1): 57

[207] Guo J E, Hu Z L, Guo X H, Zhang L C, Yu X H, Zhou S G, Chen G P. Molecular characterization of nine tissue-specific or stress-responsive genes of histone deacetylase in tomato (Solanum lycopersicum). Journal of Plant Growth Regulation, 2017, 36(3): 566–577

[208] Guo J E, Hu Z, Zhu M, Li F, Zhu Z, Lu Y, Chen G. The tomato histone deacetylase SlHDA1 contributes to the repression of fruit ripening and carotenoid accumulation. Scientific Reports, 2017, 7(1): 7930

[209] Guo J E, Hu Z, Li F, Zhang L, Yu X, Tang B, Chen G. Silencing of histone deacetylase SlHDT3 delays fruit ripening and suppresses carotenoid accumulation in tomato. Plant Science, 2017, 265: 29–38

[210] Zhu Z, An F, Feng Y, Li P, Xue L, A M, Jiang Z, Kim J M, To T K, Li W, Zhang X, Yu Q, Dong Z, Chen W Q, Seki M, Zhou J M, Guo H. Derepression of ethylene-stabilized transcription factors (EIN3/EIL1) mediates jasmonate and ethylene signaling synergy in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(30): 12539–12544

[211] Pi L, Aichinger E, van der Graaff E, Llavata-Peris C I, Weijers D, Hennig L, Groot E, Laux T. Organizer-derived WOX5 signal maintains root columella stem cells through chromatin-mediated repression of CDF4 expression. Developmental Cell, 2015, 33(5): 576–588

[212] 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. Banana transcription factor MaERF11 recruits histone deacetylase MaHDA1 and represses the expression of MaACO1 and expansins during fruit ripening. Plant Physiology, 2016, 171(2): 1070–1084

[213] Velasco R, Zharkikh A, Affourtit J, Dhingra A, Cestaro A, Kalyanaraman A, Fontana P, Bhatnagar S K, Troggio M, Pruss D, Salvi S, Pindo M, Baldi P, Castelletti S, Cavaiuolo M, Coppola G, Costa F, Cova V, Dal Ri A, Goremykin V, Komjanc M, Longhi S, Magnago P, Malacarne G, Malnoy M, Micheletti D, Moretto M, Perazzolli M, Si-Ammour A, Vezzulli S, Zini E, Eldredge G, Fitzgerald L M, Gutin N, Lanchbury J, Macalma T, Mitchell J T, Reid J, Wardell B, Kodira C, Chen Z, Desany B, Niazi F, Palmer M, Koepke T, Jiwan D, Schaeffer S, Krishnan V, Wu C, Chu V T, King S T, Vick J, Tao Q, Mraz A, Stormo A, Stormo K, Bogden R, Ederle D, Stella A, Vecchietti A, Kater M M, Masiero S, Lasserre P, Lespinasse Y, Allan A C, Bus V, Chagné D, Crowhurst R N, Gleave A P, Lavezzo E, Fawcett J A, Proost S, Rouzé P, Sterck L, Toppo S, Lazzari B, Hellens R P, Durel C E, Gutin A, Bumgarner R E, Gardiner S E, Skolnick M, Egholm M, Van de Peer Y, Salamini F, Viola R. The genome of the domesticated apple (Malus × domestica Borkh.). Nature Genetics, 2010, 42(10): 833–839

[214] D’Hont A, Denoeud F, Aury J M, Baurens F C, Carreel F, Garsmeur O, Noel B, Bocs S, Droc G, Rouard M, Da Silva C, Jabbari K, Cardi C, Poulain J, Souquet M, Labadie K, Jourda C, Lengellé J, Rodier-Goud M, Alberti A, Bernard M, Correa M, Ayyampalayam S, Mckain M R, Leebens-Mack J, Burgess D, Freeling M, Mbéguié-A-Mbéguié D, Chabannes M, Wicker T, Panaud O, Barbosa J, Hribova E, Heslop-Harrison P, Habas R, Rivallan R, Francois P, Poiron C, Kilian A, Burthia D, Jenny C, Bakry F, Brown S, Guignon V, Kema G, Dita M, Waalwijk C, Joseph S, Dievart A, Jaillon O, Leclercq J, Argout X, Lyons E, Almeida A, Jeridi M, Dolezel J, Roux N, Risterucci A M, Weissenbach J, Ruiz M, Glaszmann J C, Quétier F, Yahiaoui N, Wincker P. The banana (Musa acuminata) genome and the evolution of monocotyledonous plants. Nature Genetics, 2012, 488(7410): 213–217

[215] Garcia-Mas J, Benjak A, Sanseverino W, Bourgeois M, Mir G, González V M, Hénaff E, Câmara F, Cozzuto L, Lowy E, Alioto T, Capella-Gutiérrez S, Blanca J, Cañizares J, Ziarsolo P, Gonzalez-Ibeas D, Rodríguez-Moreno L, Droege M, Du L, Alvarez-Tejado M, Lorente-Galdos B, Melé M, Yang L, Weng Y, Navarro A, Marques-Bonet T, Aranda M A, Nuez F, Picó B, Gabaldón T, Roma G, Guigó R, Casacuberta J M, Arús P, Puigdomènech P. The genome of melon (Cucumis melo L.). Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(29): 11872–11877

[216] Ming R, Hou S, Feng Y, Yu Q, Dionne-Laporte A, Saw J H, Senin P, Wang W, Ly B V, Lewis K L, Salzberg S L, Feng L, Jones M R, Skelton R L, Murray J E, Chen C, Qian W, Shen J, Du P, Eustice M, Tong E, Tang H, Lyons E, Paull R E, Michael T P, Wall K, Rice D W, Albert H, Wang M L, Zhu Y J, Schatz M, Nagarajan N, Acob R A, Guan P, Blas A, Wai C M, Ackerman C M, Ren Y, Liu C, Wang J, Wang J, Na J K, Shakirov E V, Haas B, Thimmapuram J, Nelson D, Wang X, Bowers J E, Gschwend A R, Delcher A L, Singh R, Suzuki J Y, Tripathi S, Neupane K, Wei H, Irikura B, Paidi M, Jiang N, Zhang W, Presting G, Windsor A, Navajas-Pérez R, Torres M J, Feltus F A, Porter B, Li Y, Burroughs A M, Luo M C, Liu L, Christopher D A, Mount S M, Moore P H, Sugimura T, Jiang J, Schuler M A, Friedman V, Mitchell-Olds T, Shippen D E, dePamphilis C W, Palmer J D, Freeling M, Paterson A H, Gonsalves D, Wang L, Alam M. The draft genome of the transgenic tropical fruit tree papaya (Carica papaya Linnaeus). Nature Genetics, 2008, 452(7190): 991–996

[217] Wu J, Wang Z, Shi Z, Zhang S, Ming R, Zhu S, Khan M A, Tao S, Korban S S, Wang H, Chen N J, Nishio T, Xu X, Cong L, Qi K, Huang X, Wang Y, Zhao X, Wu J, Deng C, Gou C, Zhou W, Yin H, Qin G, Sha Y, Tao Y, Chen H, Yang Y, Song Y, Zhan D, Wang J, Li L, Dai M, Gu C, Wang Y, Shi D, Wang X, Zhang H, Zeng L, Zheng D, Wang C, Chen M, Wang G, Xie L, Sovero V, Sha S, Huang W, Zhang S, Zhang M, Sun J, Xu L, Li Y, Liu X, Li Q, Shen J, Wang J, Paull R E, Bennetzen J L, Wang J, Zhang S. The genome of the pear (Pyrus bretschneideri Rehd.). Genome Research, 2013, 23(2): 396–408

[218] Fei Z, Tang X, Alba R, Giovannoni J. Tomato Expression Database (TED): a suite of data presentation and analysis tools. Nucleic Acids Research, 2006, 34(Suppl_1): D766–D770

Acknowledgements

This work was supported by the National Natural Science Foundation of China (31722047) and the Liaoning Revitalization Talents Program (XLYC1802019).

Compliance with ethics guidelines

Yinglin Ji, Mingyang Xu, and Aide Wang declare that they have no conflicts of interest or financial conflicts to disclose. This article does not contain any studies with human or animal subjects performed by any of the authors.

版权

2021 The Author(s) 2021. Published by Higher Education Press. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0)