Molecular mechanisms involved in the cold tolerance response and regulation of fruits and vegetables

Sijie Wang , Chunru Yin , Yin Yuan , Miaomiao He , Olubukola Oluranti Babalola , Dan Qiu , Wei Deng

Horticulture Advances ›› 2026, Vol. 4 ›› Issue (1) : 4

PDF
Horticulture Advances ›› 2026, Vol. 4 ›› Issue (1) :4 DOI: 10.1007/s44281-025-00093-4
Review
review-article

Molecular mechanisms involved in the cold tolerance response and regulation of fruits and vegetables

Author information +
History +
PDF

Abstract

Fresh fruits and vegetables are critical sources of essential nutrients and natural pigments, playing a significant role in human health. However, low-temperature stress represents a major abiotic factor influencing plant growth and development. Exposure to low temperatures during the growing phase can markedly diminish both fruit yield and quality. Additionally, postharvest handling, including transportation, retail, and storage, accelerates senescence and spoilage, resulting in considerable economic losses. Although cold storage effectively reduces respiration rates and prolongs shelf life, improper application can lead to chilling injury in cold-sensitive produce, further exacerbating commercial losses. Chilling injury impairs hormone balance, disrupts cellular membrane integrity, damages photosynthetic function, and alters enzyme activity. This review examines the mechanisms underlying chilling injury and cold resistance in produce. Focusing on recent advances in cold tolerance research, particularly using Arabidopsis thaliana as a model system, it discusses the latest insights into chilling injury. Additionally, the physiological foundation of cold resistance and the role of plant hormones in this process are explored. The conclusion synthesizes identified research gaps, highlights enduring challenges, and proposes directions for future research.

Keywords

Fruits and vegetables / Chilling injury / CBF / Plant hormones

Cite this article

Download citation ▾
Sijie Wang, Chunru Yin, Yin Yuan, Miaomiao He, Olubukola Oluranti Babalola, Dan Qiu, Wei Deng. Molecular mechanisms involved in the cold tolerance response and regulation of fruits and vegetables. Horticulture Advances, 2026, 4(1): 4 DOI:10.1007/s44281-025-00093-4

登录浏览全文

4963

注册一个新账户 忘记密码

References

[1]

Aghdam MS, Luo Z, Jannatizadeh A, Sheikh-Assadi M, Sharafi Y, Farmani Bet al.. Employing exogenous melatonin applying confers chilling tolerance in tomato fruits by upregulating ZAT2/6/12 giving rise to promoting endogenous polyamines, proline, and nitric oxide accumulation by triggering arginine pathway activity.. Food Chem, 2019, 275: 549-556

[2]

An JP, Yao JF, Wang XN, You CX, Wang XF, Hao YJ. MdHY5 positively regulates cold tolerance via CBF-dependent and CBF-independent pathways in apple. J Plant Physiol, 2017, 218: 275-281

[3]

An JP, Li R, Qu FJ, You CX, Wang XF, Hao YJ. An apple NAC transcription factor negatively regulates cold tolerance via CBF-dependent pathway. J Plant Physiol, 2018, 221: 74-80

[4]

An JP, Wang XF, Zhang XW, Xu HF, Bi SQ, You CXet al.. An apple MYB transcription factor regulates cold tolerance and anthocyanin accumulation and undergoes MIEL1-mediated degradation. Plant Biotechnol J, 2020, 18: 337-353

[5]

An JP, Wang XF, Zhang XW, You CX, Hao YJ. Apple B-box protein BBX37 regulates jasmonic acid mediated cold tolerance through the JAZ-BBX37-ICE1-CBF pathway and undergoes MIEL1-mediated ubiquitination and degradation. New Phytol, 2021, 229: 2707-2729

[6]

An JP, Xu RR, Liu X, Su L, Yang K, Wang XFet al.. Abscisic acid insensitive 4 interacts with ICE1 and JAZ proteins to regulate ABA signaling-mediated cold tolerance in apple. J Exp Bot, 2022, 73: 980-997

[7]

Bai L, Liu Y, Mu Y, Anwar A, He C, Yan Y, et al. Heterotrimeric G-Protein gamma subunit CsGG3.2 positively regulates the expression of CBF genes and chilling tolerance in cucumber. Front Plant Sci. 2018;9:488. https://doi.org/10.3389/fpls.2018.00488.

[8]

Bao H, Yuan L, Luo Y, Zhang J, Liu X, Wu Qet al.. The transcription factor WRKY41-flavonoid 3'-hydroxylase module fine-tunes flavonoid metabolism and cold tolerance in potato. Plant Physiol, 2025, 197 kiaf070

[9]

Cao K, Zhang S, Chen Y, Ye J, Wei Y, Jiang Set al.. ERF transcription factor PpRAP2.12 activates PpVIN2 expression in peach fruit and reduces tolerance to cold stress. Postharvest Biol Technol, 2023, 199 112276

[10]

Che L, Lu S, Gou H, Li M, Guo L, Yang Jet al.. VvJAZ13 positively regulates cold tolerance in Arabidopsis and grape. Int J Mol Sci, 2024, 25 4458

[11]

Chen J, Li Y, Li F, Wu Q, Jiang Y, Yuan D. Banana MaABI5 is involved in ABA-induced cold tolerance through interaction with a RING E3 ubiquitin ligase, MaC3HC4-1. Sci Hortic, 2018, 237: 239-246

[12]

Chen Y, Chen L, Sun X, Kou S, Liu T, Dong Jet al.. The mitogen-activated protein kinase kinase MKK2 positively regulates constitutive cold resistance in the potato. Environ Exp Bot, 2022, 194 104702

[13]

Chen Y, Sun J, Wei Y, Cao K, Jiang S, Shao X. PpZAT10 negatively regulates peach cold resistance predominantly mediated by enhancing VIN activity. Postharvest Biol Technol, 2022, 190 111952

[14]

Chen S, Liao N, Bi H, Xu L, Wang Y, Mao Bet al.. RsWRKY49 promotes cold tolerance via activating the expression of RsCBF2 and RsNR2 in radish ( Raphanus sativus L.). Plant J, 2025, 122 e70256

[15]

Chen S, Xu L, Wang Y, Mao B, Zhang X, Song Qet al.. RsWRKY40 coordinates the cold stress response by integrating RsSPS1-mediated sucrose accumulation and the CBF-dependent pathway in radish (Raphanus sativus L.). Mol Hortic, 2025, 5 14

[16]

Ding Y, Shi Y, Yang S. Advances and challenges in uncovering cold tolerance regulatory mechanisms in plants. New Phytol, 2019, 222: 1690-1704

[17]

Ding Y, Shi Y, Yang S. Molecular regulation of plant responses to environmental temperatures. Mol Plant, 2020, 13: 544-564

[18]

Ding Y, Shi Y, Yang S. Regulatory networks underlying plant responses and adaptation to cold stress. Annu Rev Genet, 2024, 58: 43-65

[19]

Dong Y, Tang M, Huang Z, Song J, Xu J, Ahammed GJet al.. The miR164a-NAM3 module confers cold tolerance by inducing ethylene production in tomato. Plant J, 2022, 111: 440-456

[20]

Fang P, Wang Y, Wang M, Wang F, Chi C, Zhou Yet al.. Crosstalk between brassinosteroid and redox signaling contributes to the activation of CBF expression during cold responses in tomato. Antioxidants, 2021, 10 509

[21]

Feng XM, Zhao Q, Zhao LL, Qiao Y, Xie XB, Li HFet al.. The cold-induced basic helix-loop-helix transcription factor gene MdCIbHLH1 encodes an ICE-like protein in apple. BMC Plant Biol, 2012, 12 22

[22]

Fowler S, Thomashow MF. Arabidopsis transcriptome profiling indicates that multiple regulatory pathways are activated during cold acclimation in addition to the CBF cold response pathway. Plant Cell, 2002, 14: 1675-1690

[23]

Gaschler MM, Stockwell BR. Lipid peroxidation in cell death. Biochem Biophys Res Commun, 2017, 482: 419-425

[24]

Geng J, Liu JH. The transcription factor CsbHLH18 of sweet orange functions in modulation of cold tolerance and homeostasis of reactive oxygen species by regulating the antioxidant gene. J Exp Bot, 2018, 69: 2677-2692

[25]

Guo X, Liu D, Chong K. Cold signaling in plants: insights into mechanisms and regulation. J Integr Plant Biol, 2018, 60: 745-756

[26]

Guo Y, Li J, Liu L, Liu J, Li C, Yuan Let al.. The Ca2+ channels CNGC2 and CNGC20 mediate methyl jasmonate-induced calcium signaling and cold tolerance. Plant Physiol, 2025, 198 kiaf219

[27]

Gusain S, Joshi S, Joshi R. Sensing, signalling, and regulatory mechanism of cold-stress tolerance in plants. Plant Physiol Biochem, 2023, 197 107646

[28]

Han YC, Fu CC. Cold-inducible MaC2H2s are associated with cold stress response of banana fruit via regulating MaICE1. Plant Cell Rep, 2019, 38: 673-680

[29]

Han J, Li X, Li W, Yang Q, Li Z, Cheng Zet al.. Isolation and preliminary functional analysis of FvICE1, involved in cold and drought tolerance in Fragaria vesca through overexpression and CRISPR/Cas9 technologies. Plant Physiol Biochem, 2023, 196: 270-280

[30]

He M, Zhang X, Ma Y, Zhang X, Chen S, Zhu Yet al.. RsCDF3, a member of Cycling Dof Factors, positively regulates cold tolerance via auto-regulation and repressing two RsRbohs transcription in radish ( Raphanus sativus L.). Plant Sci, 2023, 337 111880

[31]

Hou XM, Zhang HF, Liu SY, Wang XK, Zhang YM, Meng YCet al.. The NAC transcription factor CaNAC064 is a regulator of cold stress tolerance in peppers. Plant Sci, 2020, 291 110346

[32]

Hou Y, Wang L, Zhao L, Xie B, Hu S, Chen Get al.. CaCl2 mitigates chilling injury in loquat fruit via the CAMTA5-mediated transcriptional repression of membrane lipid degradation genes. Food Res Int, 2022, 162 111966

[33]

Hou Y, Liu Y, Zhao L, Zhao Y, Wu Z, Zheng Yet al.. EjCML19 and EjWRKY7 synergistically function in calcium chloride-alleviated chilling injury of loquat fruit. Postharvest Biol Technol, 2023, 203 112417

[34]

Hu Y, Jiang L, Wang F, Yu D. Jasmonate regulates the inducer of CBF expression-C-repeat binding factor/DRE binding factor1 cascade and freezing tolerance in Arabidopsis. Plant Cell, 2013, 25: 2907-2924

[35]

Hu S, Wang T, Shao Z, Meng F, Chen H, Wang Qet al.. Brassinosteroid biosynthetic gene SlCYP90B3 alleviates chilling injury of tomato (Solanum lycopersicum) fruits during cold storage. Antioxidants, 2022, 11 115

[36]

Hu Y, Zhang H, Gu B, Zhang J. The transcription factor VaMYC2 from Chinese wild Vitis amurensis enhances cold tolerance of grape (V. vinifera) by up-regulating VaCBF1 and VaP5CS. Plant Physiol Biochem, 2022, 192: 218-229

[37]

Huang J, Zhao X, Bürger M, Chory J, Wang X. The role of ethylene in plant temperature stress response. Trends Plant Sci, 2023, 28: 808-824

[38]

Hwarari D, Guan Y, Ahmad B, Movahedi A, Min T, Hao Zet al.. ICE-CBF-COR signaling cascade and its regulation in plants responding to cold stress. Int J Mol Sci, 2022, 23 1549

[39]

Iqbal Z, Shariq Iqbal M, Singh SP, Buaboocha T. Ca2+/calmodulin complex triggers CAMTA transcriptional machinery under stress in plants: signaling cascade and molecular regulation. Front Plant Sci, 2020, 11 598327

[40]

Jia Z, Bao Y, Zhao Y, Liu Y, Zheng Y, Feng Zet al.. Cold shock treatment enhances cold tolerance in peach fruit through modulating PpbZIP9 and PpVIP1-mediated respiratory metabolism. Postharvest Biol Technol, 2023, 204 112421

[41]

Jiang H, Zhou LJ, Gao HN, Wang XF, Li ZW, Li YY. The transcription factor MdMYB2 influences cold tolerance and anthocyanin accumulation by activating SUMO E3 ligase MdSIZ1 in apple. Plant Physiol, 2022, 189: 2044-2060

[42]

Li H, Ye K, Shi Y, Cheng J, Zhang X, Yang S. BZR1 positively regulates freezing tolerance viaCBF-dependent and CBF-independent pathways in Arabidopsis. Mol Plant, 2017, 10: 545-559

[43]

Li C, Sun Y, Li J, Zhang T, Zhou F, Song Qet al.. ScCBF1 plays a stronger role in cold, salt and drought tolerance than StCBF1 in potato (Solanum tuberosum). J Plant Physiol, 2022, 278 153806

[44]

Li B, Wang X, Wang X, Xi Z. An AP2/ERF transcription factor VvERF63 positively regulates cold tolerance in Arabidopsis and grape leaves. Environ Exp Bot, 2023, 205 105124

[45]

Li W, Wei Y, Zhang L, Wang Y, Song P, Li Xet al.. FvMYB44, a strawberry R2R3-MYB transcription factor, improved salt and cold stress tolerance in transgenic Arabidopsis. Agronomy, 2023, 13 1051

[46]

Li B, Zang Y, Song C, Wang X, Wu X, Wang Xet al.. VvERF117 positively regulates grape cold tolerance through direct regulation of the antioxidative gene BAS1. Int J Biol Macromol, 2024, 268 131804

[47]

Liu X, Liu B, Xue S, Cai Y, Qi W, Jian Cet al.. Cucumber (Cucumis sativus L.) nitric oxide synthase associated gene1 (CsNOA1) plays a role in chilling stress. Front Plant Sci, 2016, 7 1652

[48]

Liu Y, Shi Y, Zhu N, Zhong S, Bouzayen M, Li Z. SlGRAS4 mediates a novel regulatory pathway promoting chilling tolerance in tomato. Plant Biotechnol J, 2020, 18: 1620-1633

[49]

Liu G, Zhang Z, Tian Y, Yang J, Xu X, Liu X. VvbZIP22 regulates quercetin synthesis to enhance cold resistance in grape. Plant Sci, 2025, 350 112293

[50]

Liu ZY, Qiao ZW, Ren YX, An JP, Xu SZ, Gu KDet al.. MYB20, an R2R3-type MYB transcription factor, negatively regulates salt and cold stress tolerance in pears. Plant Physiol Biochem, 2025, 229 110065

[51]

Luo H, Guan Y, Zhang Z, Zhang Z, Zhang Z, Li H. FveDREB1B improves cold tolerance of woodland strawberry by positively regulating FveSCL23 and FveCHS. Plant Cell Environ, 2024, 47: 4630-4650

[52]

Luo X, Ye X, Chen M, Zhao D, Li F. Comprehensive transcriptome analysis reveals StMAPK7 regulates cold response in potato. Plant Physiol Biochem, 2025, 223 109743

[53]

Lv K, Xie Y, Yu Q, Zhang N, Zheng Q, Wu Jet al.. Amur grape VaMYB4a-VaERF054-like module regulates cold tolerance through a regulatory feedback loop. Plant Cell Environ, 2025, 48: 1130-1148

[54]

Ma NN, Zuo YQ, Liang XQ, Yin B, Wang GD, Meng QW. The multiple stress-responsive transcription factor SlNAC1 improves the chilling tolerance of tomato. Physiol Plant, 2013, 149: 474-486

[55]

Ma X, Chen C, Yang M, Dong X, Lv W, Meng Q. Cold-regulated protein (SlCOR413IM1) confers chilling stress tolerance in tomato plants. Plant Physiol Biochem, 2018, 124: 29-39

[56]

Ma X, Gai WX, Li Y, Yu YN, Ali M, Gong ZH. The CBL-interacting protein kinase CaCIPK13 positively regulates defence mechanisms against cold stress in pepper. J Exp Bot, 2022, 73: 1655-1667

[57]

Ma X, Yu YN, Jia JH, Li QH, Gong ZH. The pepper MYB transcription factor CaMYB306 accelerates fruit coloration and negatively regulates cold resistance. Sci Hortic, 2022, 295 110892

[58]

Ma S, Lin Q, Wu T, Chen H, Hu S, Wu Bet al.. EjCBF3 conferred cold-resistance through the enhancement of antioxidase activity in loquat (Eriobotrya japonica Lindl.). Sci Hortic, 2024, 337 113556

[59]

Manasa SL, Panigrahy M, Panigrahi KCS, Rout GR. Overview of cold stress regulation in plants. Bot Rev, 2021, 88: 359-387

[60]

Mei C, Yang J, Mei Q, Jia D, Yan P, Feng Bet al.. Mdnac104 positively regulates apple cold tolerance via CBF-dependent and CBF-independent pathways. Plant Biotechnol J, 2023, 21: 2057-2073

[61]

Meng D, Li S, Feng X, Di Q, Zhou M, Yu Xet al.. CsBPC2 is essential for cucumber survival under cold stress. BMC Plant Biol, 2023, 23 566

[62]

Min D, Li F, Zhang X, Cui X, Shu P, Dong Let al.. SlMYC2 involved in methyl jasmonate-induced tomato fruit chilling tolerance. J Agric Food Chem, 2018, 66: 3110-3117

[63]

Min D, Zhou J, Li J, Ai W, Li Z, Zhang Xet al.. SlMYC2 targeted regulation of polyamines biosynthesis contributes to methyl jasmonate-induced chilling tolerance in tomato fruit. Postharvest Biol Technol, 2021, 174 111443

[64]

Nian Y, Aslam MM, Wang X, Gu H, Li W, Shao Y. The CpCOR1 gene enhances cold tolerance and antioxidant activity of papaya fruit in response to postharvest chilling stress. Postharvest Biol Technol, 2024, 218 113154

[65]

Nolan TM, Vukašinović N, Liu D, Russinova E, Yin Y. Brassinosteroids: multidimensional regulators of plant growth, development, and stress responses. Plant Cell, 2020, 32(2): 295-318

[66]

Pareek A, Khurana A, Sharma AK, Kumar R. An overview of signaling regulons during cold stress tolerance in plants. Curr Genomics, 2017, 18: 498-511

[67]

Park SU, Jung YJ, Kwon HJ, Lee JY, An J, Lee HUet al.. IbMPK3/IbMPK6-mediated IbSPF1 phosphorylation promotes cold stress tolerance in sweet potato. Biochem Biophys Res Commun, 2025, 769 151893

[68]

Provart NJ, Gil P, Chen W, Han B, Chang HS, Wang Xet al.. Gene expression phenotypes of Arabidopsis associated with sensitivity to low temperatures. Plant Physiol, 2003, 132: 893-906

[69]

Qin H, Cui X, Shu X, Zhang J. The transcription factor VaNAC72-regulated expression of the VaCP17 gene from Chinese wild Vitis amurensis enhances cold tolerance in transgenic grape (V. vinifera). Plant Physiol Biochem, 2023, 200 107768

[70]

Raza A, Charagh S, Najafi-Kakavand S, Abbas S, Shoaib Y, Anwar Set al.. Role of phytohormones in regulating cold stress tolerance: physiological and molecular approaches for developing cold-smart crop plants. Plant Stress, 2023, 8 100152

[71]

Ritonga FN, Chen S. Physiological and molecular mechanism involved in cold stress tolerance in plants. Plants, 2020, 9 560

[72]

Ritonga FN, Ngatia JN, Wang Y, Khoso MA, Farooq U, Chen S. AP2/ERF, an important cold stress-related transcription factor family in plants: A review. Physiol Mol Biol Plants, 2021, 27: 1953-1968

[73]

Sevillano L, Sanchez-Ballesta MT, Romojaro F, Flores FB. Physiological, hormonal and molecular mechanisms regulating chilling injury in horticultural species. Postharvest technologies applied to reduce its impact. J Sci Food Agric, 2009, 89: 555-573

[74]

Shan W, Kuang JF, Lu WJ, Chen JY. Banana fruit NAC transcription factor MaNAC1 is a direct target of MaICE1 and involved in cold stress through interacting with MaCBF1. Plant Cell Environ, 2014, 37: 2116-2127

[75]

Shan Y, Zhang D, Luo Z, Li T, Qu H, Duan Xet al.. Advances in chilling injury of postharvest fruit and vegetable: Extracellular ATP aspects. Compr Rev Food Sci Food Saf, 2022, 21: 4251-4273

[76]

Shi Y, Tian S, Hou L, Huang X, Zhang X, Guo Het al.. Ethylene signaling negatively regulates freezing tolerance by repressing expression of CBF and type-A ARR genes in Arabidopsis. Plant Cell, 2012, 24: 2578-2595

[77]

Song Q, Wang X, Liu Y, Brestic M, Yang X. StLTO1, a lumen thiol oxidoreductase in Solanum tuberosum L., enhances the cold resistance of potato plants. Plant Sci, 2022, 325 111481

[78]

Song J, Lin R, Tang M, Wang L, Fan P, Xia Xet al.. SlMPK1- and SlMPK2-mediated SlBBX17 phosphorylation positively regulates CBF-dependent cold tolerance in tomato. New Phytol, 2023, 239: 1887-1902

[79]

Sun X, Zhao T, Gan S, Ren X, Fang L, Karungo SKet al.. Ethylene positively regulates cold tolerance in grapevine by modulating the expression of ETHYLENE RESPONSE FACTOR 057. Sci Rep, 2016, 6 24066

[80]

Sun HJ, Luo ML, Zhou X, Zhou Q, Sun YY, Ge WYet al.. PuMYB21/PuMYB54 coordinate to activate PuPLDbeta1 transcription during peel browning of cold-stored "Nanguo" pears. Hortic Res, 2020, 7: 136

[81]

Tillett RL, Wheatley MD, Tattersall EA, Schlauch KA, Cramer GR, Cushman JC. The Vitis vinifera C-repeat binding protein 4 (VvCBF4) transcriptional factor enhances freezing tolerance in wine grape. Plant Biotechnol J, 2012, 10: 105-124

[82]

Valenzuela JL, Manzano S, Palma F, Carvajal F, Garrido D, Jamilena M. Oxidative stress associated with chilling injury in immature fruit: postharvest technological and biotechnological solutions. Int J Mol Sci, 2017, 18 1467

[83]

Wang L, Zhao R, Zheng Y, Chen L, Li R, Ma Jet al.. SlMAPK1/2/3 and antioxidant enzymes are associated with H2O2-induced chilling tolerance in tomato plants. J Agric Food Chem, 2017, 65: 6812-6820

[84]

Wang GD, Liu Q, Shang XT, Chen C, Xu N, Guan Jet al.. Overexpression of transcription factor SlNAC35 enhances the chilling tolerance of transgenic tomato. Biol Plant, 2018, 62: 479-488

[85]

Wang Y, Xu H, Liu W, Wang N, Qu C, Jiang Set al.. Methyl jasmonate enhances apple’ cold tolerance through the JAZ–MYC2 pathway. Plant Cell Tissue Organ Cult, 2018, 136: 75-84

[86]

Wang F, Chen X, Dong S, Jiang X, Wang L, Yu Jet al.. Crosstalk of PIF4 and DELLA modulates CBF transcript and hormone homeostasis in cold response in tomato. Plant Biotechnol J, 2020, 18: 1041-1055

[87]

Wang Z, Zhang Y, Hu H, Chen L, Zhang H, Chen R. CabHLH79 acts upstream of CaNAC035 to regulate cold stress in pepper. Int J Mol Sci, 2022, 23 2537

[88]

Wang T, Ma X, Chen Y, Wang C, Xia Z, Liu Zet al.. Slnac3 suppresses cold tolerance in tomatoes by enhancing ethylene biosynthesis. Plant Cell Environ, 2024, 47: 3132-3146

[89]

Wang L, Wang Y, Mao J, Zhou H, Wang L, Dai Bet al.. Methyl jasmonate regulates the glycolytic pathway and the PpbZIP43-PpFAD2 module to enhance cold resistance in peach fruit. Postharvest Biol Technol, 2025, 228 113634

[90]

Wang L, Zhao M, Zhang X, Zhao T, Huang C, Tang Yet al.. The ubiquitin ligase VviPUB19 negatively regulates grape cold tolerance by affecting the stability of ICEs and CBFs. Hortic Res, 2025, 12 uhae297

[91]

Wang X, Chen Y, Wei Y, Jiang S, Ye J, Chen Jet al.. The PpERF4/PpERF061-PpCBF1 molecular cascade regulates sucrose metabolism to influence the cold resistance of peach fruit. Postharvest Biol Technol, 2025, 230 113819

[92]

Wei Y, Li Z, Lv L, Yang Q, Cheng Z, Zhang Jet al.. Overexpression of MbICE3 increased the tolerance to cold and drought in lettuce (Lactuca sativa L.). In Vitro Cell Dev Biol Plant, 2023, 59: 767-782

[93]

Wu Y, Lv S, Zhao Y, Chang C, Hong W, Jiang J. SlHSP17.7 ameliorates chilling stress-induced damage by regulating phosphatidylglycerol metabolism and calcium signal in tomato plants. Plants, 2022, 11 1865

[94]

Xia C, Liang G, Chong K, Xu Y. The COG1-OsSERL2 complex senses cold to trigger signaling network for chilling tolerance in japonica rice. Nat Commun, 2023, 14 3104

[95]

Xiao J, Wang D, Liang L, Xie M, Tang Y, Lai YSet al.. CaMYB80 enhances the cold tolerance of pepper by directly targeting CaPOA1. Hortic Res, 2024, 11 uhae219

[96]

Xiao J, Cao B, Tang W, Sui X, Tang Y, Lai Yet al.. The CaCAD1-CaPOA1 module positively regulates pepper resistance to cold stress by increasing lignin accumulation. Int J Biol Macromol, 2025, 290 139979

[97]

Xiao J, Sui X, Xu Z, Liang L, Tang W, Tang Yet al.. CaNAC76 enhances lignin content and cold resistance in pepper by regulating CaCAD1. Int J Biol Macromol, 2025, 285 138271

[98]

Xie Y, Chen P, Yan Y, Bao C, Li X, Wang Let al.. An atypical R2R3 MYB transcription factor increases cold hardiness by CBF-dependent and CBF-independent pathways in apple. New Phytol, 2018, 218: 201-218

[99]

Xu H, Wang N, Liu J, Qu C, Wang Y, Jiang Set al.. The molecular mechanism underlying anthocyanin metabolism in apple using the MdMYB16 and MdbHLH33 genes. Plant Mol Biol, 2017, 94: 149-165

[100]

Xu XX, Hu Q, Yang WN, Jin Y. The roles of cell wall invertase inhibitor in regulating chilling tolerance in tomato. BMC Plant Biol, 2017, 17 195

[101]

Xu H, Wang N, Wang Y, Jiang S, Fang H, Zhang Jet al.. Overexpression of the transcription factor MdbHLH33 increases cold tolerance of transgenic apple callus. Plant Cell Tissue Organ Cult, 2018, 134: 131-140

[102]

Xu X, Yang H, Suo X, Liu M, Jing D, Zhang Yet al.. EjFAD8 enhances the low-temperature tolerance of loquat by desaturation of sulfoquinovosyl diacylglycerol (SQDG). Int J Mol Sci, 2023, 24: 6946

[103]

Xu J, Liu S, Hong J, Lin R, Xia X, Yu Jet al.. SlBTB19 interacts with SlWRKY2 to suppress cold tolerance in tomato via the CBF pathway. Plant J, 2024, 120: 1112-1124

[104]

Yang J, Guo X, Mei Q, Qiu L, Chen P, Li Wet al.. MdbHLH4 negatively regulates apple cold tolerance by inhibiting MdCBF1/3 expression and promoting MdCAX3L-2 expression. Plant Physiol, 2023, 191: 789-806

[105]

Ye K, Li H, Ding Y, Shi Y, Song C, Gong Zet al.. BRASSINOSTEROID-INSENSITIVE2 negatively regulates the stability of transcription factor ICE1 in response to cold stress in Arabidopsis. Plant Cell, 2019, 31: 2682-2696

[106]

Yin Q, Qin W, Zhou Z, Wu AM, Deng W, Li Zet al.. Banana MaNAC1 activates secondary cell wall cellulose biosynthesis to enhance chilling resistance in fruit. Plant Biotechnol J, 2024, 22: 413-426

[107]

Yu XH, Juan JX, Gao ZL, Zhang Y, Li WY, Jiang XM. Cloning and transformation of INDUCER of CBF EXPRESSION1 (ICE1) in tomato. Genet Mol Res, 2015, 14: 13131-13143

[108]

Yu W, Sheng J, Zhao R, Wang Q, Ma P, Shen L. Ethylene biosynthesis is involved in regulating chilling tolerance and SlCBF1 gene expression in tomato fruit. Postharvest Biol Technol, 2019, 149: 139-147

[109]

Yu W, Ma P, Sheng J, Shen L. Postharvest fruit quality of tomatoes influenced by an ethylene signaling component during long-term cold storage. Food Chem, 2023, 422 136087

[110]

Zhang W, Jiang W. UV treatment improved the quality of postharvest fruits and vegetables by inducing resistance. Trends Food Sci Technol, 2019, 92: 71-80

[111]

Zhang B, Tieman DM, Jiao C, Xu Y, Chen K, Fei Zet al.. Chilling-induced tomato flavor loss is associated with altered volatile synthesis and transient changes in DNA methylation. Proc Natl Acad Sci USA, 2016, 113: 12580-12585

[112]

Zhang L, Jiang X, Liu Q, Ahammed GJ, Lin R, Wang Let al.. The HY5 and MYB15 transcription factors positively regulate cold tolerance in tomato via the CBF pathway. Plant Cell Environ, 2020, 43: 2712-2726

[113]

Zhang W, Jiang H, Cao J, Jiang W. Advances in biochemical mechanisms and control technologies to treat chilling injury in postharvest fruits and vegetables. Trends Food Sci Technol, 2021, 113: 355-365

[114]

Zhang X, Fu X, Liu F, Wang Y, Bi H, Ai X. Hydrogen sulfide improves the cold stress resistance through the CsARF5-CsDREB3 module in cucumber. Int J Mol Sci, 2021, 22 13229

[115]

Zhang L, Song J, Lin R, Tang M, Shao S, Yu Jet al.. Tomato SlMYB15 transcription factor targeted by sly-miR156e-3p positively regulates ABA-mediated cold tolerance. J Exp Bot, 2022, 73: 7538-7551

[116]

Zhang S, Sun H, Wang J, Shen J, He F, Chen Det al.. The regulatory mechanisms and control technologies of chilling injury and fungal diseases of postharvest loquat fruit. Plants, 2022, 11 3472

[117]

Zhang Y, Peng Y, Liu J, Yan J, Zhu K, Sun Xet al.. Tetratricopeptide repeat protein SlREC2 positively regulates cold tolerance in tomato. Plant Physiol, 2023, 192: 648-665

[118]

Zhang Y, Wu L, Liu L, Jia B, Ye Z, Tang Xet al.. Functional analysis of PbbZIP11 transcription factor in response to cold stress in Arabidopsis and pear. Plants, 2023, 13 24

[119]

Zhang H, Chen M, Luo X, Song L, Li F. Overexpression of StBBX14 enhances cold tolerance in potato. Plants, 2024, 14 18

[120]

Zhang H, Guo J, Chen X, Zhou Y, Pei Y, Chen Let al.. Transcription factor CabHLH035 promotes cold resistance and homeostasis of reactive oxygen species in pepper. Hortic Plant J, 2024, 10: 823-836

[121]

Zhang L, Xing L, Dai J, Li Z, Zhang A, Wang Tet al.. Overexpression of a grape WRKY transcription factor VhWRKY44 improves the resistance to cold and salt of Arabidopsis thaliana. Int J Mol Sci, 2024, 25 7437

[122]

Zhang J, An H, Li S, Zhou B, Zhang X. Ethylene response factor EjERF23 from loquat promotes cold tolerance via directly regulating EjPOD gene participated in scavenging of hydrogen peroxide. Plant Cell Environ, 2025, 48: 7123-7138

[123]

Zhao ML, Wang JN, Shan W, Fan JG, Kuang JF, Wu KQet al.. Induction of jasmonate signalling regulators MaMYC2s and their physical interactions with MaICE1 in methyl jasmonate-induced chilling tolerance in banana fruit. Plant Cell Environ, 2012, 36: 30-51

[124]

Zhao K, Chen R, Duan W, Meng L, Song H, Wang Qet al.. Chilling injury of tomato fruit was alleviated under low-temperature storage by silencing Sly-miR171e with short tandem target mimic technology. Front Nutr, 2022, 9 906227

[125]

Zhao L, Zhao Y, Wang L, Hou Y, Bao Y, Jia Zet al.. Hot water treatment improves peach fruit cold resistance through PpHSFA4c-mediated HSF-HSP and ROS pathways. Postharvest Biol Technol, 2023, 199 112272

[126]

Zheng Q, Yu Q, Wu N, Yao W, Li J, Lv Ket al.. A grape VvHOS1-interacting HIPP protein (VvHIPP21) negatively regulates cold and drought stress. Environ Exp Bot, 2023, 207 105203

[127]

Zhu W, Li H, Dong P, Ni X, Fan M, Yang Yet al.. Low temperature-induced regulatory network rewiring via WRKY regulators during banana peel browning. Plant Physiol, 2023, 193: 855-873

[128]

Zhu Y, Zhu G, Xu R, Jiao Z, Yang J, Lin Tet al.. A natural promoter variation of SlBBX31 confers enhanced cold tolerance during tomato domestication. Plant Biotechnol J, 2023, 21: 1033-1043

[129]

Zhu X, Tang C, Zhang T, Zhang S, Wu J, Wang P. PbrCSP1, a pollen tube-specific cold shock domain protein, is essential for the growth and cold resistance of pear pollen tubes. Mol Breed, 2024, 44 18

Funding

National Key Research and Development Program of China(2022YFD2100103)

National Natural Science Foundation of China(32172596)

the Fundamental Research Funds for the Central Universities(2024CDJGF-011)

RIGHTS & PERMISSIONS

The Author(s)

PDF

4

Accesses

0

Citation

Detail

Sections
Recommended

/