果实采后冷害及其品质劣变研究进展

doi:10.16420/j.issn.0513-353x.2026-0388

摘要/Abstract

摘要：

冷害是制约冷敏性果实低温贮藏期间品质保持的关键瓶颈之一。本文中聚焦冷害诱导的果实品质劣变，系统阐述其在生物学表征、色泽劣变、香气丧失、风味失衡及质地异常等方面的变化规律，并从细胞学变化、代谢重塑、转录级联调控、表观遗传修饰、翻译后调控及信号转导网络等层面，梳理了果实冷害及其品质劣变的分子调控机制。在此基础上，结合模式植物抗冷研究的最新进展，对果实采后抗冷与品质劣变控制机制解析及保鲜技术研发进行了展望，以期为果实冷害防控及品质精准调控提供理论参考。

关键词: 果实, 采后, 冷害, 品质劣变, 色泽, 芳香, 风味, 质地, 调控机制

Abstract:

Chilling injury constitutes a major constraint on quality maintainance of chilling sensitive fruit during cold storage. This review focuses on chilling injury-induced quality deterioration in fruits，specifically addressing the associated biological manifestations，including color deterioration，aroma loss，flavor imbalance，and textural disorders. The underlying molecular regulatory networks are systematically examined from the perspectives of cellular changes，metabolic remodelling，transcriptional cascades，epigenetic modifications，post-translational regulation and signal transduction pathways. Based on recent advances in cold tolerance mechanisms from model plants，strategies for dissecting chilling resistance in postharvest fruits and developing targeted preservation technologies are further discussed. This review provides a theoretical basis for mitigating chilling injury and achieving precision regulation of fruit quality.

Key words: fruit, postharvest, chilling injury, quality deterioration, color, aroma, flavor, texture, regulatory mechanism

刘梦斐, 李俊铎, 胡若倩, 石艳娜, 徐昌杰, 陈昆松. 果实采后冷害及其品质劣变研究进展[J]. 园艺学报, 2026, 53(7): 1871-1890.

LIU Mengfei, LI Junduo, HU Ruoqian, SHI Yanna, XU Changjie, CHEN Kunsong. Advances in Chilling Injury and Associated Quality Deterioration in Postharvest Fruits[J]. Acta Horticulturae Sinica, 2026, 53(7): 1871-1890.

图/表 4

参考文献 90

[91]	Xu M, Li S J, Liu X F, Yin X R, Grierson D, Chen K S. 2019b. Ternary complex EjbHLH1-EjMYB2-EjAP2- 1 retards low temperature-induced flesh lignification in loquat fruit. Plant Physiology and Biochemistry,139:731-737.
[92]	Xu M, Zhang M X, Shi Y N, Liu X F, Li X, Grierson D, Chen K S. 2019a. EjHAT1 participates in heat alleviation of loquat fruit lignification by suppressing the promoter activity of key lignin monomer synthesis gene EjCAD5. Journal of Agricultural and Food Chemistry, 67 (18):5204-5211. doi: 10.1021/acs.jafc.9b00641 URL
[93]	Xu Q, Wang W Q, Zeng J K, Zhang J, Grierson D, Li X, Yin X R, Chen K S. 2015. A NAC transcription factor,EjNAC1,affects lignification of loquat fruit by regulating lignin. Postharvest Biology and Technology,102:25-31.
[94]	Xu Q, Yin X R, Zeng J K, Ge H, Song M, Xu C J, Li X, Ian B. Ferguson, Chen K S. 2014. Activator- and repressor-type MYB transcription factors are involved in chilling injury induced flesh lignification in loquat via their interactions with the phenylpropanoid pathway. Journal of Experimental Botany, 65 (15):4349-4359. doi: 10.1093/jxb/eru208 pmid: 24860186
[95]	Yang C, Duan W Y, Xie K L, Ren C H, Zhu C Q, Chen K S, Zhang B. 2020. Effect of salicylic acid treatment on sensory quality,flavor-related chemicals and gene expression in peach fruit after cold storage. Postharvest Biology and Technology, 161:111089. doi: 10.1016/j.postharvbio.2019.111089 URL
[96]	Yang S L, Sun C D, Wang P, Shan L L, Cai C, Zhang B, Zhang W S, Li X, Ferguson I, Chen K S. 2008. Expression of expansin genes during postharvest lignification and softening of ‘Luoyangqing’ and ‘Baisha’ loquat fruit under different storage conditions. Postharvest Biology and Technology,49:46-53.
[1]	Albornoz K, Zhou J Q, Yu J W, Beckles D M. 2022. Dissecting postharvest chilling injury through biotechnology. Current Opinion in Biotechnology, 78:102790. doi: 10.1016/j.copbio.2022.102790 URL
[2]	Ai S J, Liang L, Liu M F, Grierson D, Chen K S, Xu C J. 2025. PpERF17 alleviates peach fruit postharvest chilling injury under elevated CO₂ by activating jasmonic acid and γ-aminobutyric acid biosynthesis. Horticulture Research,12:uhaf014.
[3]	Bai C M, Fang M H, Zhai B Q, Ma L L, Fu A Z, Gao L P, Kou X H, Meng D M, Wang Q, Zheng S F, Zuo J H. 2021. Regulations of m⁶A methylation on tomato fruit chilling injury. Horticultural Plant Journal,7:434-442.
[4]	Bao Y Q, Xie B, Hou Y Y, Zhao L Y, Hu S Q, Zhao Y Q, Zheng Y H, Wang P, Jin P. 2024a. PpCaM7 and PpCAMTA5 synergistically mitigate chilling-induced browning by suppressing reactive oxygen species synthesis in peach fruit. Postharvest Biology and Technology, 214:112978. doi: 10.1016/j.postharvbio.2024.112978 URL
[5]	Bao Y Q, Zeng Y, Zhao L Y, Liu Y, Zhao Y Q, Xiao L H, Wu Z G, Zheng Y H, Jin P. 2025. Phytosulfokine α treatment induces osmotic regulation to alleviate postharvest chilling injury of peach fruit by PpbHLH62. Postharvest Biology and Technology, 230:113746. doi: 10.1016/j.postharvbio.2025.113746 URL
[6]	Bao Z Y, Zhou Q H, Yu Y, Chen W, Yang Z F, Cao S F, Shi L Y. 2024b. Melatonin treatment induces DNA methylation to alleviate chilling induced-browning in cold stored peach fruit. Postharvest Biology and Technology, 208:112686. doi: 10.1016/j.postharvbio.2023.112686 URL
[97]	Ye Y J, Xiao Y Y, Han Y C, Shan W, Fan Z Q, Xu Q G, Kuang J F, Lu W J, Lakshmanan P, Chen J Y. 2016. Banana fruit VQ motif-containing protein5 represses cold-responsive transcription factor MaWRKY26 involved in the regulation of JA biosynthetic genes. Scientific Reports, 6:23632. doi: 10.1038/srep23632
[98]	Yin Q, Qin W Q, Zhou Z B, Wu A M, Deng W, Li Z G, Shan W, Chen J Y, Kuang J F, Lu W J. 2024. Banana MaNAC1 activates secondary cell wall cellulose biosynthesis to enhance chilling resistance in fruit. Plant Biotechnology Journal,22:413-426.
[99]	Yu W Q, Jia X X, Shen L, Sheng J P. 2023. Ethylene perception influences the chilling injury and SlCBF1 gene expression during cold storage of tomato fruit. Postharvest Biology and Technology, 204:112434. doi: 10.1016/j.postharvbio.2023.112434 URL
[100]	Zeng J K, Li X, Xu Q, Chen J Y, Yin X R, Ferguson I B, Chen K S. 2015. EjAP2-1,an AP2/ERF gene,is a novel regulator of fruit lignification induced by chilling injury,via interaction with EjMYB transcription factors. Plant Biotechnology Journal, 13 (9):1325-1334. doi: 10.1111/pbi.2015.13.issue-9 URL
[101]	Zhang B, Tieman D M, Jiao C, Xu Y M, Chen K S, Fei Z J, Giovannoni J J, Klee H J. 2016. Chilling-induced tomato flavor loss is associated with altered volatile synthesis and transient changes in DNA methylation. Proceedings of the National Academy of Sciences of the United States of America, 113 (4):12580-12585.
[102]	Zhang B, Xi W P, Wei W W, Shen J Y, Ferguson I, Chen K S. 2011. Changes in aroma-related volatiles and gene expression during low temperature storage and subsequent shelf-life of peach fruit. Postharvest Biology and Technology,60:7-16.
[7]	Calixto C P G, Guo W B, James A B, Tzioutziou N A, Entizne J C, Panter P E, Knight H, Nimmo H G, Zhang R X, Brown J W S. 2018. Rapid and dynamic alternative splicing impacts the Arabidopsis cold response transcriptome. The Plant Cell,30:1424-1444.
[8]	Cantín C M, Crisosto C H, Ogundiwin E A, Gradziel T, Torrents J, Moreno M A, Gogorcena Y. 2010. Chilling injury susceptibility in an intra-specific peach [Prunus persica(L.) Batsch] progeny. Postharvest Biology and Technology,58:79-87.
[9]	Carvajal F, Palma F, Jamilena M, Garrido D. 2015. Cell wall metabolism and chilling injury during postharvest cold storage in zucchini fruit. Postharvest Biology and Technology,108:68-77.
[10]	Chen G, Kuang X Y, Fan Z Q, Chen Y Z, Lin Y F, Wang H, Chen Y H, Lin H T. 2024. Energy and proline metabolisms participate in the occurrence of chilling injury in cold-stored Chinese olive fruit. Postharvest Biology and Technology, 214:112955. doi: 10.1016/j.postharvbio.2024.112955 URL
[11]	Chen Y N, Lin S Q, Song W, Wei W, Shan W, Kuang J F, Chen J Y, Lu W J. 2025. MaWRKY21 is involved in ethylene-mediated cold tolerance in banana fruit by regulating ROS accumulation. Postharvest Biology and Technology, 226:113541. doi: 10.1016/j.postharvbio.2025.113541 URL
[12]	Dawson D M, Melton L D, Watkins C B. 1992. Cell wall changes in nectarines(Prunus persica):solubilization and depolymerization of pectic and neutral polymers during ripening and in mealy fruit. Plant Physiology, 100 (3):1203-1210. doi: 10.1104/pp.100.3.1203 pmid: 16653106
[103]	Zhang B, Zhang S Y, Wu Y J, Li Y, Kong L Y, Wu R R, Zhao M, Liu W, Yu H. 2024a. Defining context-dependent m⁶A RNA methylomes in Arabidopsis. Developmental Cell, 59 (20):2772-2786.e3. doi: 10.1016/j.devcel.2024.06.012 URL
[104]	Zhang G H, Zhang Y, Kou X H, Li J K, Luo D L, Huang T Y, Wang X F, Cao S. 2024b. Salicylic acid mitigates chilling injury to peaches by improving antioxidant capacity and energy metabolism. Scientia Horticulturae, 338:113841. doi: 10.1016/j.scienta.2024.113841 URL
[105]	Zhang J, Yin X R, Li H, Xu M, Zhang M X, Li S J, Liu X F, Shi Y N, Grierson D, Chen K S. 2020. ETHYLENE RESPONSE FACTOR39-MYB8 complex regulates low-temperature-induced lignification of loquat fruit. Journal of Experimental Botany, 71 (10):3172-3184. doi: 10.1093/jxb/eraa085 pmid: 32072171
[106]	Zhang J, Zhang M X, Xu M, Yin X R, Grierson D, Chen K S. 2018. EjMYB4 is a transcriptional activator of 4-coumarate:coenzyme A ligase involved in lignin biosynthesis in loquat(Eriobotrya japonica). Plant Growth Regulation,86:413-421.
[107]	Zhang M X, Shi Y N, Liu Z M, Zhang Y J, Yin X R, Liang Z H, Huang Y Q, Grierson D, Chen K S. 2022a. An EjbHLH14-EjHB1-EjPRX12 module is involved in methyl jasmonate alleviation of chilling-induced lignin deposition in loquat fruit. Journal of Experimental Botany, 73 (5):1668-1682. doi: 10.1093/jxb/erab511 URL
[13]	Ding Y L, Shi Y T, Yang S H. 2024. Regulatory networks underlying plant responses and adaptation to cold stress. Annual Review of Genetics,58:43-65.
[14]	Ding Y L, Yang H, Wu S F, Fu D Y, Li M Z, Gong Z Z, Yang S H. 2022. CPK28-NLP 7 module integrates cold-induced Ca²⁺ signal and transcriptional reprogramming in Arabidopsis. Science Advances, 8 (26):eabn7901.
[15]	Dixon R A, Barros J. 2019. Lignin biosynthesis:old roads revisited and new roads explored. Open Biol, 9:190215. doi: 10.1098/rsob.190215 URL
[16]	Duan W Y, Yang C, Cao X M, Wei C Y, Chen K S, Li X, Zhang B. 2023. Chilling-induced peach flavor loss is associated with expression and DNA methylation of functional genes. Journal of Advanced Research,53:17-31.
[17]	Duan W Y, Yang C, Cao X M, Zhang C, Liu H R, Chen K S, Li X, Zhang B. 2022. Transcriptome and DNA methylome analysis reveal new insights into methyl jasmonate-alleviated chilling injury of peach fruit after cold storage. Postharvest Biology and Technology, 189:111915. doi: 10.1016/j.postharvbio.2022.111915 URL
[18]	Fan Z Q, Chen J Y, Kuang J F, Lu W J, Shan W. 2017. The banana fruit SINA ubiquitin ligase MaSINA1 regulates the stability of MaICE1 to be negatively involved in cold stress response. Frontiers in Plant Science, 8:995. doi: 10.3389/fpls.2017.00995 URL
[19]	Farneti B, Alarcón A A, Papasotiriou F G, Samudrala D, Cristescu S M, Costa G, Harren F J M, Woltering E J. 2015. Chilling-induced changes in aroma volatile profiles in tomato. Food and Bioprocess Technology,8:1442-1454.
[108]	Zhang M Z, Zhang Q, Tian C, Liu G S, Pan Y G, Xu X B, Shi X Q, Zhang Z K, Meng L H. 2022b. Physiological and transcriptome analyses of CaCl₂ treatment to alleviate chilling injury in pineapple. Plants, 11 (16):2215. doi: 10.3390/plants11172215 URL
[109]	Zhang S Y, Cao K F, Wei Y Y, Jiang S, Ye J F, Xu F, Chen Y, Shao X F. 2023. PpBZR1,a BES/BZR transcription factor,enhances cold stress tolerance by suppressing sucrose degradation in peach fruit. Plant Physiology and Biochemistry, 202:107972. doi: 10.1016/j.plaphy.2023.107972 URL
[110]	Zhang W F, Gong Z H, Wu M B, Chan Helen, Yuan Y J, Tang N, Zhang Q, Miao M J, Chang W, Li Z, Li Z G, Jin L, Deng W. 2019. Integrative comparative analyses of metabolite and transcript profiles uncovers complex regulatory network in tomato(Solanum lycopersicum L.)fruit undergoing chilling injury. Scientific Reports, 9:4470. doi: 10.1038/s41598-019-41065-9
[111]	Zhao K Y, Chen R L, Duan W H, Meng L H, Song H M, Wang Q, Li J K, Xu X B. 2022. Chilling injury of tomato fruit was alleviated under low-temperature storage by silencing Sly-miR171e with short tandem target mimic technology. Frontiers in Nutrition, 9:2022.
[112]	Zhao L Y, Cassan-Wang H, Zhao Y Q, Bao Y Q, Hou Y Y, Liu Y, Wu Z G, Bouzayen M, Zheng Y H, Jin P. 2025a. Calcium-dependent protein kinase PpCDPK29-mediated Ca²⁺-ROS signal and PpHSFA2a phosphorylation regulate postharvest chilling tolerance of peach fruit. Plant Biotechnology Journal,23:1938-1953.
[20]	Fu X D, Li F J, Li Y N, Li X A, Zhang X H, Ahmed Z F R. 2025. New insights into controlling horticultural crop quality deterioration caused by cold stress:epigenetic modification. Horticulture Research, 13 (3):uhaf326.
[21]	Ge H, Zhang J, Zhang Y J, Li X, Yin X R, Grierson D, Chen K S. 2017. EjNAC3 transcriptionally regulates chilling-induced lignification of loquat fruit via physical interaction with an atypical CAD-like gene. Journal of Experimental Botany, 68 (18):5129-5136. doi: 10.1093/jxb/erx330 URL
[22]	Gross K C, Wang C Y, Saltveit M E. 2016. The commercial storage of fruits,vegetables,and florist and nursery stocks. Beltsville: United States Department of Agriculture,Agricultural Research Service
[23]	Guillén F, Medina-Santamarina J, García-Pastor M E, Chen N J, Uruu G, Paull R E. 2022. Postharvest melatonin treatment delays senescence and increases chilling tolerance in pineapple. LWT-Food Science and Technology, 169:113989. doi: 10.1016/j.lwt.2022.113989 URL
[24]	He X X, Wei Y Y, Kou J Y, Xu F, Chen Z H, Shao X F. 2018. PpVIN2,an acid invertase gene family member,is sensitive to chilling temperature and affects sucrose metabolism in postharvest peach fruit. Plant Growth Regulation,86:169-180.
[25]	He Y, Li W H, Yang W T, Dai M, Shi J Y, Song Z Y. 2025. PpZAT6-PpWRKY 46 module regulates chilling stress response of peach fruit by activating proline biosynthesis. Postharvest Biology and Technology, 230:113733. doi: 10.1016/j.postharvbio.2025.113733 URL
[26]	Hou Y Y, Li Z Y, Zheng Y H, Jin P. 2021. Effects of CaCl₂ treatment alleviates chilling injury of loquat fruit(Eriobotrya japonica)by modulating ROS homeostasis. Foods, 10 (7):1662. doi: 10.3390/foods10071662 URL
[27]	Hou Y Y, Wang L, Zhao L Y, Xie B, Hu S Q, Chen G F, Zheng Y H, Jin P. 2022a. CaCl₂ mitigates chilling injury in loquat fruit via the CAMTA5-mediated transcriptional repression of membrane lipid degradation genes. Food Research International, 162:111966. doi: 10.1016/j.foodres.2022.111966 URL
[28]	Hou Y Y, Zhao L Y, Xie B, Hu S Q, Zheng Y H, Jin P. 2022b. EjCaM7 and EjCAMTA3 synergistically alleviate chilling-induced lignification in loquat fruit by repressing the expression of lignin biosynthesis genes. Postharvest Biology and Technology, 192:112010. doi: 10.1016/j.postharvbio.2022.112010 URL
[29]	Huang D, Xue L, Lu Y Q, Liu M F, Lin-Wang K, Allan A C, Zhang B, Chen K S, Xu C J. 2024a. PpBBX32 and PpZAT5 modulate temperature-dependent and tissue-specific anthocyanin accumulation in peach fruit. Horticulture Research,11:uhae212.
[30]	Huang H, Jian Q J, Jiang Y M, Duan X W, Qu H X. 2016. Enhanced chilling tolerance of banana fruit treated with malic acid prior to low-temperature storage. Postharvest Biology and Technology,111:209-213.
[31]	Huang W N, Shi Y N, Yan H, Wang H, Wu D, Grierson D, Chen K S. 2023. The calcium-mediated homogalacturonan pectin complexation in cell walls contributes the firmness increase in loquat fruit during postharvest storage. Journal of Advanced Research,49:47-62.
[32]	Huang Y Q, Liang Z H, Lu J, Zhang M X, Cao X Z, Hu R Q, Li D D, Grierson D, Chen W B, Zhu C Q, Wu D, Shi Y N, Chen K S. 2024b. The transcription factor EjNAC 5 regulates loquat fruit chilling lignification. Journal of Experimental Botany, 75 (20):6625-6643. doi: 10.1093/jxb/erae333 URL
[33]	Jiao C F. 2022. PpCBF6 is involved in phytosulfokine α-retarded chilling injury by suppressing the expression of PpLOX5 in peach fruit. Frontiers in Plant Science, 13:874338. doi: 10.3389/fpls.2022.874338 URL
[34]	Jiao C F, Wei Y C, Dong J. 2023. PpMYB105 inhibits chilling injury by regulating PpMsrA1 in peach fruit. Plant Cell Reports, 42 (10):1557-1569. doi: 10.1007/s00299-023-03047-0
[35]	Jiao Jianqing. 2022. Regulatory mechanism of postharvest lignification induced by low temperature in kiwifruit[Ph. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	焦健青. 2022. 低温诱导猕猴桃果实采后木质化调控机理研究[博士论文]. 杨凌: 西北农林科技大学.
[36]	Ketsa S, Chidtragool S, Klein J D, Lurie S. 1999. Firmness,pectin components and cell wall hydrolases of mango fruit following low temperature stress. The Journal of Horticultural Science and Biotechnology, 74 (6):685-689. doi: 10.1080/14620316.1999.11511173 URL
[37]	Kindgren P, Ard R, Ivanov M, Marquardt S. 2018. Transcriptional read-through of the long non-coding RNA SVALKA governs plant cold acclimation. Nature Communications, 9:4561. doi: 10.1038/s41467-018-07010-6 pmid: 30385760
[38]	Kong X M, Zhou Q, Zhou X, Wei B D, Ji S J. 2020. Transcription factor CaNAC1 regulates low-temperature-induced phospholipid degradation in green bell pepper. Journal of Experimental Botany, 71 (3):1078-1091.
[39]	Li X, Tian Y J, Duan X, Lv M Y, Shao X F, Wei Y Y, Mason P, Guan R N, Wang Q Y, Pang X Y, Zhang G Q, Henry R J. 2026. Single-cell and spatial transcriptomics reveals the spatiotemporal trajectory of the small peptide TAP4 in delaying postharvest fruit senescence. Plant Communications,doi:10.1016/j.xplc.2026.101846.
[40]	Liang Z H, Shi Y N, Huang Y Q, Lu J, Zhang M X, Cao X Z, Hu R Q, Li D D, Chen W B, Zhu C Q, Wu D, Chen K S. 2024. Xylem NAC domain 1(EjXND1)relieves cold-induced lignification by negatively regulating the EjHB1-EjPRX12 module in loquat fruit. Journal of Advanced Research,73:93-104.
[113]	Zhao L Y, Jia Z Y, Zhang Y, Bao Y Q, Zhao Y Q, Liu Y, Xi Y Y, Zheng Y H, Jin P. 2026a. Heat treatment-induced PpbZIP43 regulates energy metabolism to enhance postharvest cold tolerance in peach fruit. Postharvest Biology and Technology, 233:114032. doi: 10.1016/j.postharvbio.2025.114032 URL
[114]	Zhao M L, Wang J N, Shan W, Fan J G, Kuang J F, Wu K Q, Li X P, Chen W X, He F Y, Chen J Y, Lu W J. 2013. Induction of jasmonate signalling regulators MaMYC2s and their physical interactions with MaICE1 in methyl jasmonate-induced chilling tolerance in banana fruit. Plant,Cell and Environment,36:30-51.
[115]	Zhao Y Q, Liu Z A, Liu Y, Bao Y Q, Zhao L Y, Wu Z G, Zheng Y H, Jin P. 2025b. PpCML30-PpWRKY11 module regulated PpOAT expression and proline accumulation in H₂S-induced cold tolerance in peach fruit. Postharvest Biology and Technology, 226:113544. doi: 10.1016/j.postharvbio.2025.113544 URL
[116]	Zhao Y Q, Liu Z A, Zhao L Y, Xi Y Y, Wu Z G, Zheng Y H, Jin P. 2026b. Transcription factor PpMYB73 inhibits PpPPO 1 transcription and flesh browning in hydrogen sulfide-mitigated chilling injury of peach fruit. Postharvest Biology and Technology, 232:114004. doi: 10.1016/j.postharvbio.2025.114004 URL
[117]	Zhao Y Y, Bao X, Bi J F. 2026c. ABA mitigates chilling injury in peach fruit via the PpFERL-mediated sucrose metabolism. International Journal of Biological Macromolecules, 338:149720. doi: 10.1016/j.ijbiomac.2025.149720 URL
[118]	Zheng Y H, Chen X T, Chen W, Yang Z F, Li X W, Cao S F, Shi L Y. 2025. Inhibition of DNA methylation modification regulates endogenous GABA,proline and AsA-GSH metabolism and enhances antioxidant capacity to improve chilling tolerance in postharvest kiwifruit. Postharvest Biology and Technology, 230:113812. doi: 10.1016/j.postharvbio.2025.113812 URL
[41]	Lin H, Bai L J, Wei W, Su W B, Wu Y T, Wu R, Wang H, Chen J Y, Lin H T, Fan Z Q. 2024a. The role of MaWRKY70 in regulating lipoxygenase gene transcription during chilling injury development in banana fruit. Foods, 13:854. doi: 10.3390/foods13060854 URL
[42]	Lin S Q, Song W, Gan L Z, Wei W, Shan W, Kuang J F, Chen J Y, Lu W J. 2024b. Low temperature downregulates MabHLH355 and its associated target genes responsible for scavenging ROS in banana peel under cold stress. Postharvest Biology and Technology, 213:112956. doi: 10.1016/j.postharvbio.2024.112956 URL
[43]	Lin Z X, Cai D L, Liu Z L, Lin Z X, Wei W, Shan W, Chen J Y, Lu W J, Su X G, Kuang J F. 2025. MaWRKY45 mediates H₂S-mitigated chilling injury in banana fruit by coordinating with MaERF53L/121L. Postharvest Biology and Technology, 230:113749. doi: 10.1016/j.postharvbio.2025.113749 URL
[44]	Liu H R, He H, Liu C X, Wang C F, Qiao Y J, Zhang B. 2022. Changes of sensory quality,flavor-related metabolites and gene expression in peach fruit treated by controlled atmosphere(CA)under cold storage. International Journal of Molecular Sciences, 23:7141. doi: 10.3390/ijms23137141 URL
[45]	Liu H R, He H, Wang Q K, Li S Z, Guo F J, Zhang C F, Wang K, Liu C X, Wang C F, Wang X, Chen B J, Zhang Y, Hu L S, Qiao Y J. 2025a. Near-freezing temperature alters ROS metabolism and gene transcription to enhance peach chilling tolerance and aroma quality. Food Research International, 209:116297. doi: 10.1016/j.foodres.2025.116297 URL
[46]	Liu J, Li Q X, Chen J J, Jiang Y M. 2020. Revealing further insights on chilling injury of postharvest bananas by untargeted lipidomics. Foods, 9:894. doi: 10.3390/foods9070894 URL
[47]	Liu Q B, Ding Y L, Shi Y T, Ma L, Wang Y, Song C P, Wilkins K A, Davies J M, Knight H, Knight M R, Gong Z Z, Guo Y, Yang S H. 2021. The calcium transporter ANNEXIN1 mediates cold‐induced calcium signaling and freezing tolerance in plants. The EMBO Journal,40:e104559.
[48]	Liu Y, Bao Y Q, Zhao Y Q, Zhao L Y, Liu Z A, Xiao L H, Zheng Y H, Jin P. 2026. Phytosulfokine α mitigates cold-induced lignification in loquat fruit through suppressing EjWRKY33 and EjbHLH77 module-mediated lignin biosynthesis. The EMBO Journal Postharvest Biology and Technology, 233:114065.
[49]	Liu Z A, Zhao Y Q, Liu Y, Zhao L Y, Bao Y Q, Xiang W, Cai X Y, Zheng Y H, Jin P. 2025b. Exogenous H₂S promoted osmoregulatory capability via PpWRKY11-mediated transcriptional activation of PpGAD1 in cold-stored peach fruit. Plant Physiology and Biochemistry, 228:110258. doi: 10.1016/j.plaphy.2025.110258 URL
[50]	Lurie S. 2022. Proteomic and metabolomic studies on chilling injury in peach and nectarine. Frontiers in Plant Science, 13:958312. doi: 10.3389/fpls.2022.958312 URL
[51]	Lyons J M. 1973. Chilling injury in plants. Annual Review of Plant Physiology,24:445-466.
[52]	Ma Y, Dai X Y, Xu Y Y, Luo W, Zheng X M, Zeng D L, Pan Y J, Lin X L, Liu H H, Zhang D J, Xiao J, Guo X Y, Xu S J, Niu Y D, Jin J B, Zhang H, Xu X, Li L G, Wang W, Qian Q, Ge S, Chong K. 2015. COLD 1 confers chilling tolerance in rice. Cell,160:1209-1221.
[53]	Moison M, Pacheco J M, Lucero L, Fonouni-Farde C, Rodríguez-Melo J, Mansilla N, Christ A, Bazin J, Benhamed M, Ibañez F, Crespi M, Estevez J M, Ariel F. 2021. The lncRNA APOLO interacts with the transcription factor WRKY 42 to trigger root hair cell expansion in response to cold. Molecular Plant,14:937-948.
[54]	Pang Xuequn, Chen Yanni, Huang Xuemei, Wang Yuehua, Hu Weirong, Zhang Zhaoqi. 2008. Chilling temperature enhances quality reduction in postharvest ‘Shatangju’(Citrus reticulata Blanco)fruit. Acta Horticulturae Sinica, 35 (4):509-514. (in Chinese)
	庞学群, 陈燕妮, 黄雪梅, 汪跃华, 胡位荣, 张昭其. 2008. 冷害导致砂糖橘果实品质劣变. 园艺学报, 35 (4):509-514.
[55]	Peace C P, Crisosto C H, Garner D T, Dandekar A M, Gradziel T M, Bliss F A. 2006. Genetic control of internal breakdown in peach. Acta Horticulturae,713:489-496.
[56]	Pelloux J, Rustérucci C, Mellerowicz E J. 2007. New insights into pectin methylesterase structure and function. Trends in Plant Science, 12 (6):267-277. doi: 10.1016/j.tplants.2007.04.001 pmid: 17499007
[57]	Peng C, Hua W, Liu J. 2025. Unraveling the signaling pathways of plant cold stress:current insights and future directions. Frontiers in Plant Science, 16:1666852. doi: 10.3389/fpls.2025.1666852 URL
[58]	Peng H H, Shan W, Kuang J F, Lu W J, Chen J Y. 2013. Molecular characterization of cold-responsive basic helix-loop-helix transcription factors MabHLHs that interact with MaICE1 in banana fruit. Planta,238:937-953.
[59]	Ponce-Valadez M, Escalona-Buendía H B, Villa-Hernández J M, León-Sánchez F D, Rivera-Cabrera F, Alia-Tejacal I, Pérez-Flores L J. 2016. Effect of refrigerated storage(12.5 ℃)on tomato(Solanum lycopersicum)fruit flavor:a biochemical and sensory analysis. Postharvest Biology and Technology,111:6-14.
[60]	Prasanna V, Prabha T N, Tharanathan R N. 2007. Fruit ripening phenomena - an overview. Critical Reviews in Food Science and Nutrition, 47 (1):1-19. doi: 10.1080/10408390600976841 pmid: 17364693
[61]	Rugkong A, Rose J K C, Lee S J, Giovannoni J J, O’Neill M A, Watkins C B. 2010. Cell wall metabolism in cold-stored tomato fruit. Postharvest Biology and Technology, 57 (2):106-113. doi: 10.1016/j.postharvbio.2010.03.004 URL
[62]	Sevillano L, Sanchez-Ballesta M T, Romojaro F, Flores F B. 2009. Physiological, hormonal and molecular mechanisms regulating chilling injury in horticultural species. Postharvest technologies applied to reduce its impact. Journal of the Science of Food and Agriculture,89:555-573.
[63]	Si J, Fan Z Q, Wu C J, Yang Y Y, Shan W, Kuang J F, Lu W J, Wei W, Chen J Y. 2024. MaHsf24,a novel negative modulator,regulates cold tolerance in banana fruits by repressing the expression of HSPs and antioxidant enzyme genes. Plant Biotechnology Journal,22:2873-2886.
[64]	Song F Y, Huangfu Z Q, Han Y R, Li H M, Wang Z P, Jin X W, Chen J L. 2024. Nitric oxide fumigation can affect the metabolism of volatile compounds derived from analyses of fatty acids and amino acids in post-harvest flat peach during cold storage. Food Control, 159:110258. doi: 10.1016/j.foodcont.2023.110258 URL
[65]	Song F, Li Z X, Wang C, Jiang Y C, Wang Z J, He L G, Ma X F, Zhang Y, Song X, Liu J H, Wu L M. 2023. CsMYB15 positively regulates Cs4CL2-mediated lignin biosynthesis during juice sac granulation in navel orange. Frontiers in Plant Science, 14:1223820. doi: 10.3389/fpls.2023.1223820 URL
[66]	Song C B, Wu M B, Zhou Y, Gong Z H, Yu W W, Zhang Y, Yang Z F. 2022. NAC-mediated membrane lipid remodeling negatively regulates fruit cold tolerance. Horticulture Research,9:uhac039.
[67]	Song C B, Yang Y Y, Yang T W, Ba L J, Zhang H, Han Y C, Xiao Y Y, Shan W, Kuang J F, Chen J Y, Lu W J. 2019. MaMYB4 recruits histone deacetylase MaHDA2 and modulates the expression of ω-3 fatty acid desaturase genes during cold stress response in banana fruit. Plant and Cell Physiology,60:2410-2422.
[68]	Stanley C J, Scofield C, Hallett I C, Schröder R. 2023. Dissecting the role of cell wall changes in chilling injury-induced gel formation,rubberiness,and mealiness in apricots. Horticulturae, 9 (11):1115. doi: 10.3390/horticulturae9101115 URL
[69]	Sun B L, Kuang X Y, Lin H T, Lin M S, Chen Y Z, Zeng L Z, Lin Y F, Chen Y H, Wang H, Fan Z Q. 2023. The role of respiratory metabolism in chilling injury development of Chinese olive fruit during cold storage. Postharvest Biology and Technology, 205:112489. doi: 10.1016/j.postharvbio.2023.112489 URL
[70]	Sun H J, Luo M L, Zhou X, Zhou Q, Sun Y Y, Ge W Y, Yao M M, Ji S J. 2020. PuMYB21/PuMYB54 coordinate to activate PuPLDβ1 transcription during peel browning of cold-stored‘Nanguo’pears. Horticulture Research, 7:136. doi: 10.1038/s41438-020-00356-3
[71]	Tan Xiaoyu, Niu Junpeng, Wang Quanzhi, Zhang Li, Zhu Lijuan, Wang Guodong, Zheng Heyun, Yu Zhifang, Duan Yuquan, Jiang Li, Sun Xiuxiu, Yang Ying, Luo Weiqi, Li Xuehui, Guan Le, Zhao Yanling, Li Guoxiao, Yin Congfei, Ge Cheng, Ma Min, Jia Luting, Zhang Xu, Zhao Yaoyao, Geng Xinli, Wang Libin, Zhang Shaoling. 2025. Recent advances in JAs-induced improvement of postharvest chilling resistance in horticultural crops. Acta Horticulturae Sinica, 52 (8):1987-2020. (in Chinese) doi: 10.16420/j.issn.0513-353x.2024-0969
[119]	Zhou Cheng, Fang Yi, Zhou Jinyang, Huang Qihao, Pan Yongjian, Shi Qianqian, Ni Huixian, Yang Zhenfeng, Song Chunbo. 2023. The relationship between membrane lipid metabolism and chilling injury of postharvest peach fruit induced by low temperature. Acta Horticulturae Sinica, 50 (6):1305-1317. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-0427
	周成, 方怡, 周锦杨, 黄企浩, 盘永坚, 史千千, 倪慧娴, 杨震峰, 宋春波. 2023. 低温诱导的桃果实采后膜脂代谢变化与冷害的关系. 园艺学报, 50 (6):1305-1317. doi: 10.16420/j.issn.0513-353x.2022-0427
[120]	Zhou D D, Zhang Q, Li P X, Pan L Q, Tu K. 2020. Combined transcriptomics and proteomics analysis provides insight into metabolisms of sugars,organic acids and phenols in UV-C treated peaches during storage. Plant Physiology and Biochemistry,157:148-159.
[121]	Zhou H J, Ye Z W, Wang L F, Zhang S Y, Yuan Z Y, Su M S, Zhang X A, Du J H, Li X W, Zhang M H, Hu Y, Shen S L, Zheng X L, Huan C. 2025. 1-MCP regulates taste development in cold-stored peach fruit through modulation of sugar,organic acid,and polyphenolic metabolism. Postharvest Biology and Technology, 225:113518. doi: 10.1016/j.postharvbio.2025.113518 URL
[122]	Zhou H J, Zhang X A, Su M S, Du J H, Li X W, Zhang M H, Hu Y, Huan C, Ye Z W. 2022. Controlled atmosphere storage alleviates internal browning in flat peach fruit by regulating energy and sugar metabolisms. Plant Physiology and Biochemistry,186:107-120.
[123]	Zhou H W, Lurie S, Lers A, Khatchitski A, Sonego L, Ben Arie R. 2000a. Delayed storage and controlled atmosphere storage of nectarines:two strategies to prevent woolliness. Postharvest Biology and Technology, 18 (2):133-141. doi: 10.1016/S0925-5214(99)00072-1 URL
[124]	Zhou H W, Sonego L, Khalchitski A, Ben-Arie R, Lers A, Lurie S. 2000b. Cell wall enzymes and cell wall changes in‘Flavortop’nectarines:mRNA abundance,enzyme activity,and changes in pectic and neutral polymers during ripening and in woolly fruit. Journal of the American Society for Horticultural Science, 125 (5):630-637. doi: 10.21273/JASHS.125.5.630 URL
[71]	谭筱玉, 牛军鹏, 王全智, 张丽, 朱丽娟, 王国栋, 郑贺云, 郁志芳, 段玉权, 姜丽, 孙秀秀, 杨瑛, 罗伟奇, 李雪晖, 管乐, 赵艳岭, 李国晓, 殷从飞, 葛成, 马敏, 贾璐婷, 张旭, 赵垚垚, 耿新丽, 王利斌, 张绍铃. 2025. 茉莉酸及其衍生物提高园艺作物采后低温抗性的研究进展. 园艺学报, 52 (8):1987-2020. doi: 10.16420/j.issn.0513-353x.2024-0969
[72]	Tian J X, Xie S Y, Zhang P, Wang Q, Li J K, Xu X B. 2022. Attenuation of postharvest peel browning and chilling injury of banana fruit by Astragalus polysaccharides. Postharvest Biology and Technology, 184:111783. doi: 10.1016/j.postharvbio.2021.111783 URL
[73]	Tian Yu, Pan Wencheng, Yang Jiayu, Wei Yaxiong, Zhu Yanzhe, Huang Yanfa, Wu Longguo. 2025. Comprehensive evaluation of storage and preservation effects of exogenous GABA treatments on cherry tomato. Acta Horticulturae Sinica, 52 (11):2987-3002. (in Chinese) doi: 10.16420/j.issn.0513-353x.2025-0326
	田雨, 潘文成, 杨佳玉, 魏亚雄, 朱彦哲, 黄彦发, 吴龙国. 2025. 外源GABA 处理对樱桃番茄贮藏保鲜效果综合评价. 园艺学报, 52 (11):2987-3002. doi: 10.16420/j.issn.0513-353x.2025-0326
[74]	Vicente A M, Manavski N, Rohn P T, Schmid L M, Garcia-Molina A, Leister D, Seydel C, Bellin L, Möhlmann T, Ammann G, Kaiser S, Meurer J. 2023. The plant cytosolic m⁶A RNA methylome stabilizes photosynthesis in the cold. Plant Communications, 4:100634. doi: 10.1016/j.xplc.2023.100634 URL
[75]	Wang D, Ren X Y, Meng L K, Zheng R Y, Li D, Kong Q J. 2023. Exogenous phytosulfokine α(PSKα)alleviates chilling injury of kiwifruit by regulating Ca²⁺ and protein kinase-mediated reactive oxygen species metabolism. Foods, 12:4196. doi: 10.3390/foods12234196 URL
[76]	Wang J C, Ren Yulong, Liu X, Luo S, Zhang X, Liu X, Lin Q B, Zhu S S, Wan H, Yang Y, Zhang Y, Lei B, Zhou C L, Pan T, Wang Y F, Wu M M, Ruonan jing, Xu Y, Han M, Wu F Q, Lei C L, Guo X P, Cheng Z J, Zheng X M, Wang Y H, Zhao Z J, Jiang L, Zhang X, Wang Yong-Fei, Wang H Y, Wan J M. 2021a. Transcriptional activation and phosphorylation of OsCNGC 9 confer enhanced chilling tolerance in rice. Molecular Plant, 14 (2):315-329. doi: 10.1016/j.molp.2020.11.022 URL
[125]	Zhou J X, Min D D, Li ZL, Fu X D, Zhao X M, Wang J H, Zhang X H, Li F J, Li X A. 2021. Effects of chilling acclimation and methyl jasmonate on sugar metabolism in tomato fruits during cold storage. Scientia Horticulturae, 289:110495. doi: 10.1016/j.scienta.2021.110495 URL
[126]	Zhu Y C, Wang K, Wu C X, Hao Y T, Zhang B, Grierson D, Chen K S, Xu C J. 2021. DNA hypermethylation associated with the development of temperature-dependent postharvest chilling injury in peach fruit. Postharvest Biology and Technology, 181:111645. doi: 10.1016/j.postharvbio.2021.111645 URL
[127]	Zhu Y C, Wang K, Wu C X, Zhao Y, Yin X R, Zhang B, Grierson D, Chen K S, Xu C J. 2019. Effect of ethylene on cell wall and lipid metabolism during alleviation of postharvest chilling injury in peach. Cells, 8:1612. doi: 10.3390/cells8121612 URL
[128]	Zhu Y C, Zhang B, Allan A C, Lin-Wang K, Zhao Y, Wang K, Chen K S, Xu C J. 2020. DNA demethylation is involved in the regulation of temperature-dependent anthocyanin accumulation in peach. The Plant Journal,102:965-976.
[129]	Zuo X X, Wang J, Wu Z G, Jin P, Zheng Y H. 2025. High relative humidity storage mitigates chilling injury in zucchini fruit via regulating the CpMYB63-mediated cell wall degradation. Postharvest Biology and Technology, 222:113388. doi: 10.1016/j.postharvbio.2025.113388 URL
[77]	Wang K, Yin X R, Zhang B, Grierson D, Xu C J, Chen K S. 2017. Transcriptomic and metabolic analyses provide new insights into chilling injury in peach fruit. Plant,Cell and Environment,40:1531-1551.
[78]	Wang L F, Wang Y X, Mao J X, Zhou H J, Wang L B, Dai B E, Zhou L, Zhang S Y, Shen S L, Zheng X L, Huan C. 2025a. Methyl jasmonate regulates the glycolytic pathway and the PpbZIP43-PpFAD2 module to enhance cold resistance in peach fruit. Postharvest Biology and Technology, 228:113634. doi: 10.1016/j.postharvbio.2025.113634 URL
[79]	Wang T T, Liu J C, Yuan Y B, Wang Y, Cheng C X, Yang S L. 2024. PpWRKY65 mediates peach(Prunus persica)fruit lignification in both ambient temperature storage and chilling injury condition. Postharvest Biology and Technology, 216:113043. doi: 10.1016/j.postharvbio.2024.113043 URL
[80]	Wang W Q, Zhang J, Ge H, Li S J, Li X, Yin X R, Grierson D, Chen K S. 2016. EjMYB8 transcriptionally regulates flesh lignification in loquat fruit. PLoS ONE, 11 (4):e0154399. doi: 10.1371/journal.pone.0154399 URL
[81]	Wang X X, Chen Y, Wei Y Y, Jiang S, Ye J F, Chen J H, Xu F, Elsherbiny E A, Shao X F. 2025b. The PpERF4/PpERF061-PpCBF 1 molecular cascade regulates sucrose metabolism to influence the cold resistance of peach fruit. Postharvest Biology and Technology, 230:113819. doi: 10.1016/j.postharvbio.2025.113819 URL
[82]	Wang Y X, Gao L P, Zhu B Z, Zhu H L, Luo Y B, Wang Q, Zuo J H. 2018. Integrative analysis of long non-coding RNA acting as ceRNAs involved in chilling injury in tomato fruit. Gene,667:25-33.
[83]	Wang Z Q, Pu H L, Shan S S, Zhang P, Li J K, Song H M, Xu X B. 2021b. Melatonin enhanced chilling tolerance and alleviated peel browning of banana fruit under low temperature storage. Postharvest Biology and Technology, 179:111571. doi: 10.1016/j.postharvbio.2021.111571 URL
[84]	Wasternack C, Strnad M. 2018. Jasmonates:news on occurrence,biosynthesis,metabolism and action of an ancient group of signaling compounds. International Journal of Molecular Sciences, 19:2539. doi: 10.3390/ijms19092539 URL
[85]	Wongs-Aree C, Aschariyaphotha W, Palapol Y, Bodhipadma K, Noichinda S. 2024. Structural membrane alterations in tropical horticultural crops under postharvest chilling stress. Vegetable Research,4:e016.
[86]	Wu Y Y, Zhang S Y, Chen Y, Wei Y Y, Xu F, Ding P, Shao X F. 2025. PpWRKY4 regulates chilling injury in peach fruit by activating PpVIN2 and accelerating sucrose metabolism. Postharvest Biology and Technology, 230:113729. doi: 10.1016/j.postharvbio.2025.113729 URL
[87]	Xi W P, Zhang B, Liang L, Shen J Y, Wei W W, Xu C J, Allan A C, Ferguson I B, Chen K S. 2012a. Postharvest temperature influences volatile lactone production via regulation of acyl-CoA oxidases in peach fruit. Plant,Cell and Environment, 35 (3):534-545. doi: 10.1111/pce.2012.35.issue-3 URL
[88]	Xi W P, Zhang B, Shen J Y, Sun C D, Xu C J, Chen K S. 2012b. Intermittent warming alleviated the loss of peach fruit aroma-related esters by regulation of AAT during cold storage. Postharvest Biology and Technology,74:42-48.
[89]	Xiang Miaolian, Wu Fan, Li Shucheng, Ma Qiaoli, Wang Yinbao, Xiao Liuhua, Chen Jinyin, Chen Ming. 2022. Exogenous melatonin regulates reactive oxygen metabolism to induce resistance of postharvest pear fruit to black spot. Acta Horticulturae Sinica, 49 (5):1102-1110. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0159
	向妙莲, 吴帆, 李树成, 马巧利, 王印宝, 肖刘华, 陈金印, 陈明. 2022. 外源褪黑素调控活性氧代谢诱导梨果实抗采后黑斑病. 园艺学报, 49 (5):1102-1110. doi: 10.16420/j.issn.0513-353x.2021-0159
[90]	Xiao J C, Sui X Y, Xu Z P, Liang L, Tang W, Tang Y, Sun B, Lai Y S, Huang Z, Zheng Y X, Li H X. 2025. CaNAC76 enhances lignin content and cold resistance in pepper by regulating CaCAD1. International Journal of Biological Macromolecules, 285:138271. doi: 10.1016/j.ijbiomac.2024.138271 URL

表征层次 Symptom level	主要特征与核心机制 Main characteristics and core mechanisms	关联的主要品质劣变 Associated major quality deterioration
生理生化与分子 Physiological， biochemical and molecular	蛋白质和核酸发生修饰；代谢与转运及调控因子基因表达异常，引致物质代谢全面失调；呼吸链受损，ATP合成减少，能荷下降，能量代谢障碍，物质运输受阻；ROS产生与清除失衡，氧化应激爆发，诱发氧化损伤；激素合成与信号转导异常 Protein and nucleic acid modifications；aberrant expression of metabolic，transport，and regulatory genes，leading to comprehensive metabolic disorder；impaired respiratory chain，reduced ATP synthesis，decreased energy charge，energy disturbance，and blocked transport；ROS imbalance，oxidative stress，and oxidative damage；abnormal hormone synthesis and signal transduction	成熟障碍：正常成熟进程及品质呈现全面受阻 Ripening disorder：blocked ripening process，impaired quality formation and maintenance
代谢物 Metabolite	膜脂组分重塑，不饱和脂肪酸合成受抑，膜脂饱和度上升；风味物质积累失衡，香气物质积累减少，糖酸比失调；渗透调节物质和酚类等次生代谢物质积累 Remodeling of membrane lipid composition；inhibition of unsaturated fatty acid synthesis，increased membrane lipid saturation；imbalance in flavor substance accumulation，reduced aroma compounds，altered sugar-acid ratio；abnormal accumulation of osmoregulatory substances and phenolic compounds	风味劣变：香气寡淡、异味产生、营养品质下降 Flavor deterioration：bland aroma，off-flavor production，reduced nutritional quality
细胞与组织 Cell and tissue	膜脂相变，流动性下降，膜系统崩解损伤，膜透性增加，区室化破坏，细胞与组织受损；果胶解聚受阻以及纤维素和木质素异常沉积导致细胞壁结构改变，细胞黏附异常 Membrane lipid phase transition，decreased fluidity，membrane system disruption and damage，increased membrane permeability，compartmentalization breakdown，cell and tissue damage；inhibited pectin depolymerization，abnormal deposition of cellulose and lignin，altered cell wall structure，abnormal cell adhesion	色泽劣变：褐变、水渍状 Color deterioration：browning，water soaking 质地劣变：软化或木质化 Texture deterioration：softening or lignification
外观 Appearance	果皮或果肉细胞塌陷，酚类和色素氧化导致水渍状、黑色或褐色、平整或凹陷的各种类型病斑 Collapse of peel or flesh cells；oxidation of phenolics and pigments，leading to various types of lesions including water-soaked，black or brown，flat or sunken	商品性丧失：外观差、腐烂 Loss of marketability：poor appearance，decay

表征层次 Symptom level	主要特征与核心机制 Main characteristics and core mechanisms	关联的主要品质劣变 Associated major quality deterioration
生理生化与分子 Physiological， biochemical and molecular	蛋白质和核酸发生修饰；代谢与转运及调控因子基因表达异常，引致物质代谢全面失调；呼吸链受损，ATP合成减少，能荷下降，能量代谢障碍，物质运输受阻；ROS产生与清除失衡，氧化应激爆发，诱发氧化损伤；激素合成与信号转导异常 Protein and nucleic acid modifications；aberrant expression of metabolic，transport，and regulatory genes，leading to comprehensive metabolic disorder；impaired respiratory chain，reduced ATP synthesis，decreased energy charge，energy disturbance，and blocked transport；ROS imbalance，oxidative stress，and oxidative damage；abnormal hormone synthesis and signal transduction	成熟障碍：正常成熟进程及品质呈现全面受阻 Ripening disorder：blocked ripening process，impaired quality formation and maintenance
代谢物 Metabolite	膜脂组分重塑，不饱和脂肪酸合成受抑，膜脂饱和度上升；风味物质积累失衡，香气物质积累减少，糖酸比失调；渗透调节物质和酚类等次生代谢物质积累 Remodeling of membrane lipid composition；inhibition of unsaturated fatty acid synthesis，increased membrane lipid saturation；imbalance in flavor substance accumulation，reduced aroma compounds，altered sugar-acid ratio；abnormal accumulation of osmoregulatory substances and phenolic compounds	风味劣变：香气寡淡、异味产生、营养品质下降 Flavor deterioration：bland aroma，off-flavor production，reduced nutritional quality
细胞与组织 Cell and tissue	膜脂相变，流动性下降，膜系统崩解损伤，膜透性增加，区室化破坏，细胞与组织受损；果胶解聚受阻以及纤维素和木质素异常沉积导致细胞壁结构改变，细胞黏附异常 Membrane lipid phase transition，decreased fluidity，membrane system disruption and damage，increased membrane permeability，compartmentalization breakdown，cell and tissue damage；inhibited pectin depolymerization，abnormal deposition of cellulose and lignin，altered cell wall structure，abnormal cell adhesion	色泽劣变：褐变、水渍状 Color deterioration：browning，water soaking 质地劣变：软化或木质化 Texture deterioration：softening or lignification
外观 Appearance	果皮或果肉细胞塌陷，酚类和色素氧化导致水渍状、黑色或褐色、平整或凹陷的各种类型病斑 Collapse of peel or flesh cells；oxidation of phenolics and pigments，leading to various types of lesions including water-soaked，black or brown，flat or sunken	商品性丧失：外观差、腐烂 Loss of marketability：poor appearance，decay

品质劣变 Quality deterioration	信号/处理 Signal / treatment	上游转录因子 Upstream transcription factor	末端转录因子 Terminal transcription factor	末端转录因子对靶基因及代谢的调控效应 Regulatory effects of terminal transcription factors on target genes and metabolisms	物种 Species	参考文献 Reference
色泽劣变 Color worsening	低温 Cold	—	bHLH355	激活GSTF12/T1，增强ROS清除能力，利于抗冷 Activates GSTF12/T1 expression，enhances ROS scavenging capacity，improves chilling tolerance	香蕉 Musa acuminata	Lin et al.,2024b
	热激 Heat	—	bZIP43	促进H⁺-ATPase11/SDH1，增强抗冷性 Promotes H⁺-ATPase11/SDH1，enhances chilling tolerance	桃 Prunus persica	Zhao et al.,2026a
	MeJA	—	bZIP43	激活FAD2，促进不饱和脂肪酸合成，增强膜流动性，利于抗冷 Activates FAD2，promotes unsaturated fatty acid synthesis，enhances membrane fluidity，improves chilling tolerance	桃 Prunus persica	Wang et al.,2025a
	CaCl₂	—	CAMTA5	抑制PLC6-like/LOX5，延缓膜脂降解、维持细胞膜稳定，利于抗冷 Suppresses PLC6-like/LOX5，delays membrane lipid degradation，maintains membrane stability，improves chilling tolerance	枇杷 Eriobotrya japonica	Hou et al.,2022a
	MeJA	bHLH1/2/4，MYC2a/b	CBF1/2	与ICE1互作，激活ICE-CBF冷信号通路基因，利于抗冷 Interacts with ICE1，activates ICE-CBF cold signaling pathway genes，improves chilling tolerance	香蕉 Musa acuminata	Peng et al.,2013；Zhao et al.,2013
色泽劣变 Color worsening	H₂S	WRKY45	ERF53L/121L	增强ROS清除基因表达，提升抗氧化能力，利于抗冷 Enhances the expression of ROS scavenging genes，improves antioxidant capacity，improves chilling tolerance	香蕉 Musa acuminata	Lin et al.,2025
	热激 Heat	—	HSFA2a	增强热激蛋白及抗氧化基因表达，减轻冷害 Enhances the expression of heat shock protein and antioxidant genes，alleviates chilling injury	桃 Prunus persica	Zhao et al.,2025a
	热激 Heat	—	HSF24	抑制APX/MDAR/GST，增加ROS积累，加重冷害 Suppresses APX/MDAR/GST，increases ROS accumulation，aggravates chilling injury	香蕉 Musa acuminata	Si et al.,2024
	低温 Cold	—	MYB21/54	激活PLDβ1，促进膜脂降解，果皮褐变 Activates PLDβ1，promotes membrane lipid degradation，peel browning	梨 Pyrus ussuriensis	Sun et al.,2020
	H₂S	—	MYB73	抑制PPO1，减轻冷害导致的酚类氧化 Suppresses PPO1 expression，alleviates chilling-induced phenolic oxidation	桃 Prunus persica	Zhao et al.,2026b
	MeJA	—	MYB105	激活MsrA1表达，减少ROS积累，减轻冷害 Activates MsrA1，reduces ROS accumulation，alleviates chilling injury	桃 Prunus persica	Jiao et al.,2023
	低温 Cold	—	NAC1	激活PLDα4，促进膜脂降解，破坏膜结构 Activates PLDα4，promotes membrane lipid degradation，disrupts membrane structure	辣椒 Capsicum annuum	Kong et al.,2020
	低温 Cold	CAMTA5	NAC25	激活PAO/RBOHs，促进ROS积累，加重冷害 Activates PAO/RBOHs，promotes ROS accumulation，aggravates chilling injury	桃 Prunus persica	Bao et al.,2024a
	低温 Cold	—	NAC25/28	激活PLDs/DGKs/PLCs，促进膜脂降解，降低抗冷性 Activates PLDs/DGKs/PLCs，promotes membrane lipid degradation，reduces chilling tolerance	香蕉 Musa acuminata	Song et al.,2022
	低温 Cold	—	WRKY21	激活RbohE1/H3，加剧氧化损伤 Activates RbohE1/H3，exacerbates oxidative damage	香蕉Musa acuminata	Chen et al.,2025
	MeJA	—	WRKY26	激活LOX2/AOS3/OPR3，促进JA积累，增强耐冷性 Activates LOX2/AOS3/OPR3 expression，promotes jasmonic acid accumulation，enhances chilling tolerance	香蕉 Musa acuminata	Ye et al.,2016
	低温 Cold	—	WRKY70	激活LOX1/3.1/3.2/4，加速膜脂氧化，加剧冷害 Activates LOX1/3.1/3.2/4，accelerates membrane lipid oxidation，aggravates chilling injury	香蕉 Musa acuminata	Lin et al.,2024a
风味劣变 Flavor deterioration	PSK-α	—	bHLH62	激活GAD4/P5CS/OAT表达，提升GABA、多胺及脯氨酸含量，利于抗冷 Activates GAD4/P5CS/OAT expression，increases GABA，polyamine and proline contents，improves chilling tolerance	桃 Prunus persica	Bao et al.,2025
	BR	—	BZR1	抑制VIN2，维持蔗糖含量，减轻冷害 Suppresses VIN2，maintains sucrose content，alleviates chilling injury	桃 Prunus persica	Zhang et al.,2023
	低温 Cold	ERF061/ERF4	CBF1	激活VIN2，加速蔗糖降解 Activates VIN2，accelerates sucrose degradation	桃 Prunus persica	Wang et al.,2025b
	H₂S	—	WRKY11	激活OAT，促进脯氨酸积累，利于抗冷 Activates OAT，promotes proline accumulation，improves chilling tolerance	桃 Prunus persica	Zhao et al.,2025b
	CO₂	MYC2.1	ERF17	激活LOX/AOS/OPR3/GAD，促进JA、GABA积累，增强抗冷性 Activates LOX/AOS/OPR3/GAD，promotes JA and GABA accumulation，enhances chilling tolerance	桃 Prunus persica	Ai et al.,2025
	低温 Cold	—	WRKY4	激活VIN2，促进蔗糖分解，负调控耐寒性 Activates VIN2，promotes sucrose degradation，negatively regulates chilling tolerance	桃 Prunus persica	Wu et al.,2025
	H₂S	—	WRKY11	激活GAD1，促进GABA积累，减轻冷害 Activates GAD1，promotes GABA accumulation，alleviates chilling injury	桃 Prunus persica	Liu et al.,2025b
	ABA	WRKY46	ZAT6	激活P5CS，提高脯氨酸含量，减轻冷害 Activates P5CS，increases proline content，alleviates chilling injury	桃 Prunus persica	He et al.,2025
质地劣变 Texture distortion	低温 Cold	—	bHLH1-AP2-1-MYB2	抑制4CL1，减轻木质化 Suppresses 4CL1 expression，alleviates lignification	枇杷 Eriobotrya japonica	Xu et al.,2019b
	CaCl₂	—	CAMTA3	抑制POD64/LAC12-like，减轻木质化 Suppresses POD64/LAC12-like，alleviates lignification	枇杷 Eriobotrya japonica	Hou et al.,2022b
	低温 Cold	MYB8	ERF39	激活4CL1，加重木质化 Activates 4CL1，enhances lignification	枇杷 Eriobotrya japonica	Zhang et al.,2020
	低温 Cold	NAC5	ERF39/HB1	激活4CL1/PRX12，促进木质素积累 Activates 4CL1/PRX12，promotes lignin accumulation	枇杷 Eriobotrya japonica	Huang et al.,2024b
	热激 Heat	—	HAT1	抑制CAD5，减缓冷害木质化 Suppresses CAD5，alleviates chilling-induced lignification	枇杷 Eriobotrya japonica	Xu et al.,2019a
	MeJA	bHLH14	HB1	激活PRX12，促进木质化积累 Activates PRX12，promotes lignin accumulation	枇杷 Eriobotrya japonica	Zhang et al.,2022a
	低温 Cold	XND1	HB1	激活PRX12，促进木质化 Activates PRX12，promotes lignification	枇杷 Eriobotrya japonica	Liang et al.,2024
	低温 Cold	AP2-1	MYB1/2	激活/抑制4CL1，促进/减少木质素合成 Activates/suppresses 4CL1，promotes/reduces lignin synthesis	枇杷 Eriobotrya japonica	Xu et al.,2014；Zeng et al.,2015
	低温 Cold	—	MYB4	激活4CL1，促进木质化 Activates 4CL1 expression，promotes lignification	枇杷 Eriobotrya japonica	Zhang et al.,2018
	低温 Cold	—	MYB5/12/13/ 14	激活C4H1/4CL3，促进木质化 Activates C4H1/4CL3，promotes lignification	猕猴桃 Actinidia chinensis	焦健青,2022
	低温 Cold	—	MYB8	激活4CL1，促进木质化 Activates 4CL1 expression，promotes lignification	枇杷 Eriobotrya japonica	Wang et al.,2016
	低温 Cold	—	MYB15	激活4CL2，导致木质素积累，引发汁胞粒化 Activates 4CL2 expression，leads to lignin accumulation，triggers juice sac granulation	甜橙 Citrus sinensis	Song et al.,2023
	热激 Heat	—	MYB63	激活PME3/PG1，加速细胞壁降解，加重冷害 Activates PME3/PG1，accelerates cell wall degradation，aggravates chilling injury	西葫芦 Cucurbita pepo	Zuo et al.,2025
	低温 Cold	—	NAC1	激活PAL1/4CL1，促进木质化 Activates PAL1/4CL1，promotes lignification	枇杷 Eriobotrya japonica	Xu et al.,2015
	低温 Cold	—	NAC1	激活CESA7/6B，促进细胞壁增厚，增强抗冷性 Activates CESA7/6B，promotes cell wall thickening，enhances chilling tolerance	香蕉 Musa acuminata	Yin et al.,2024
	低温 Cold	—	NAC3	激活CAD-like，促进木质素合成 Activates CAD-like，promotes lignin synthesis	枇杷 Eriobotrya japonica	Ge et al.,2017
	低温 Cold	—	NAC76	激活CAD1，增强细胞壁强度与抗冷能力 Activates CAD1，enhances cell wall strength and chilling tolerance	辣椒 Capsicum annuum	Xiao et al.,2025
	PSK-α	bHLH77	WRKY33	激活PAL/C4H/4CL/POD，促进木质化 Activates PAL/C4H/4CL/POD，promotes lignification	枇杷 Eriobotrya japonica	Liu et al.,2026
	低温 Cold	—	WRKY65	抑制4CL7/CAD1/COMT/F5H，减轻木质化 Suppresses 4CL7/CAD1/COMT/F5H expression，alleviates lignification	桃 Prunus persica	Wang et al.,2024

品质劣变 Quality deterioration	信号/处理 Signal / treatment	上游转录因子 Upstream transcription factor	末端转录因子 Terminal transcription factor	末端转录因子对靶基因及代谢的调控效应 Regulatory effects of terminal transcription factors on target genes and metabolisms	物种 Species	参考文献 Reference
色泽劣变 Color worsening	低温 Cold	—	bHLH355	激活GSTF12/T1，增强ROS清除能力，利于抗冷 Activates GSTF12/T1 expression，enhances ROS scavenging capacity，improves chilling tolerance	香蕉 Musa acuminata	Lin et al.,2024b
	热激 Heat	—	bZIP43	促进H⁺-ATPase11/SDH1，增强抗冷性 Promotes H⁺-ATPase11/SDH1，enhances chilling tolerance	桃 Prunus persica	Zhao et al.,2026a
	MeJA	—	bZIP43	激活FAD2，促进不饱和脂肪酸合成，增强膜流动性，利于抗冷 Activates FAD2，promotes unsaturated fatty acid synthesis，enhances membrane fluidity，improves chilling tolerance	桃 Prunus persica	Wang et al.,2025a
	CaCl₂	—	CAMTA5	抑制PLC6-like/LOX5，延缓膜脂降解、维持细胞膜稳定，利于抗冷 Suppresses PLC6-like/LOX5，delays membrane lipid degradation，maintains membrane stability，improves chilling tolerance	枇杷 Eriobotrya japonica	Hou et al.,2022a
	MeJA	bHLH1/2/4，MYC2a/b	CBF1/2	与ICE1互作，激活ICE-CBF冷信号通路基因，利于抗冷 Interacts with ICE1，activates ICE-CBF cold signaling pathway genes，improves chilling tolerance	香蕉 Musa acuminata	Peng et al.,2013；Zhao et al.,2013
色泽劣变 Color worsening	H₂S	WRKY45	ERF53L/121L	增强ROS清除基因表达，提升抗氧化能力，利于抗冷 Enhances the expression of ROS scavenging genes，improves antioxidant capacity，improves chilling tolerance	香蕉 Musa acuminata	Lin et al.,2025
	热激 Heat	—	HSFA2a	增强热激蛋白及抗氧化基因表达，减轻冷害 Enhances the expression of heat shock protein and antioxidant genes，alleviates chilling injury	桃 Prunus persica	Zhao et al.,2025a
	热激 Heat	—	HSF24	抑制APX/MDAR/GST，增加ROS积累，加重冷害 Suppresses APX/MDAR/GST，increases ROS accumulation，aggravates chilling injury	香蕉 Musa acuminata	Si et al.,2024
	低温 Cold	—	MYB21/54	激活PLDβ1，促进膜脂降解，果皮褐变 Activates PLDβ1，promotes membrane lipid degradation，peel browning	梨 Pyrus ussuriensis	Sun et al.,2020
	H₂S	—	MYB73	抑制PPO1，减轻冷害导致的酚类氧化 Suppresses PPO1 expression，alleviates chilling-induced phenolic oxidation	桃 Prunus persica	Zhao et al.,2026b
	MeJA	—	MYB105	激活MsrA1表达，减少ROS积累，减轻冷害 Activates MsrA1，reduces ROS accumulation，alleviates chilling injury	桃 Prunus persica	Jiao et al.,2023
	低温 Cold	—	NAC1	激活PLDα4，促进膜脂降解，破坏膜结构 Activates PLDα4，promotes membrane lipid degradation，disrupts membrane structure	辣椒 Capsicum annuum	Kong et al.,2020
	低温 Cold	CAMTA5	NAC25	激活PAO/RBOHs，促进ROS积累，加重冷害 Activates PAO/RBOHs，promotes ROS accumulation，aggravates chilling injury	桃 Prunus persica	Bao et al.,2024a
	低温 Cold	—	NAC25/28	激活PLDs/DGKs/PLCs，促进膜脂降解，降低抗冷性 Activates PLDs/DGKs/PLCs，promotes membrane lipid degradation，reduces chilling tolerance	香蕉 Musa acuminata	Song et al.,2022
	低温 Cold	—	WRKY21	激活RbohE1/H3，加剧氧化损伤 Activates RbohE1/H3，exacerbates oxidative damage	香蕉Musa acuminata	Chen et al.,2025
	MeJA	—	WRKY26	激活LOX2/AOS3/OPR3，促进JA积累，增强耐冷性 Activates LOX2/AOS3/OPR3 expression，promotes jasmonic acid accumulation，enhances chilling tolerance	香蕉 Musa acuminata	Ye et al.,2016
	低温 Cold	—	WRKY70	激活LOX1/3.1/3.2/4，加速膜脂氧化，加剧冷害 Activates LOX1/3.1/3.2/4，accelerates membrane lipid oxidation，aggravates chilling injury	香蕉 Musa acuminata	Lin et al.,2024a
风味劣变 Flavor deterioration	PSK-α	—	bHLH62	激活GAD4/P5CS/OAT表达，提升GABA、多胺及脯氨酸含量，利于抗冷 Activates GAD4/P5CS/OAT expression，increases GABA，polyamine and proline contents，improves chilling tolerance	桃 Prunus persica	Bao et al.,2025
	BR	—	BZR1	抑制VIN2，维持蔗糖含量，减轻冷害 Suppresses VIN2，maintains sucrose content，alleviates chilling injury	桃 Prunus persica	Zhang et al.,2023
	低温 Cold	ERF061/ERF4	CBF1	激活VIN2，加速蔗糖降解 Activates VIN2，accelerates sucrose degradation	桃 Prunus persica	Wang et al.,2025b
	H₂S	—	WRKY11	激活OAT，促进脯氨酸积累，利于抗冷 Activates OAT，promotes proline accumulation，improves chilling tolerance	桃 Prunus persica	Zhao et al.,2025b
	CO₂	MYC2.1	ERF17	激活LOX/AOS/OPR3/GAD，促进JA、GABA积累，增强抗冷性 Activates LOX/AOS/OPR3/GAD，promotes JA and GABA accumulation，enhances chilling tolerance	桃 Prunus persica	Ai et al.,2025
	低温 Cold	—	WRKY4	激活VIN2，促进蔗糖分解，负调控耐寒性 Activates VIN2，promotes sucrose degradation，negatively regulates chilling tolerance	桃 Prunus persica	Wu et al.,2025
	H₂S	—	WRKY11	激活GAD1，促进GABA积累，减轻冷害 Activates GAD1，promotes GABA accumulation，alleviates chilling injury	桃 Prunus persica	Liu et al.,2025b
	ABA	WRKY46	ZAT6	激活P5CS，提高脯氨酸含量，减轻冷害 Activates P5CS，increases proline content，alleviates chilling injury	桃 Prunus persica	He et al.,2025
质地劣变 Texture distortion	低温 Cold	—	bHLH1-AP2-1-MYB2	抑制4CL1，减轻木质化 Suppresses 4CL1 expression，alleviates lignification	枇杷 Eriobotrya japonica	Xu et al.,2019b
	CaCl₂	—	CAMTA3	抑制POD64/LAC12-like，减轻木质化 Suppresses POD64/LAC12-like，alleviates lignification	枇杷 Eriobotrya japonica	Hou et al.,2022b
	低温 Cold	MYB8	ERF39	激活4CL1，加重木质化 Activates 4CL1，enhances lignification	枇杷 Eriobotrya japonica	Zhang et al.,2020
	低温 Cold	NAC5	ERF39/HB1	激活4CL1/PRX12，促进木质素积累 Activates 4CL1/PRX12，promotes lignin accumulation	枇杷 Eriobotrya japonica	Huang et al.,2024b
	热激 Heat	—	HAT1	抑制CAD5，减缓冷害木质化 Suppresses CAD5，alleviates chilling-induced lignification	枇杷 Eriobotrya japonica	Xu et al.,2019a
	MeJA	bHLH14	HB1	激活PRX12，促进木质化积累 Activates PRX12，promotes lignin accumulation	枇杷 Eriobotrya japonica	Zhang et al.,2022a
	低温 Cold	XND1	HB1	激活PRX12，促进木质化 Activates PRX12，promotes lignification	枇杷 Eriobotrya japonica	Liang et al.,2024
	低温 Cold	AP2-1	MYB1/2	激活/抑制4CL1，促进/减少木质素合成 Activates/suppresses 4CL1，promotes/reduces lignin synthesis	枇杷 Eriobotrya japonica	Xu et al.,2014；Zeng et al.,2015
	低温 Cold	—	MYB4	激活4CL1，促进木质化 Activates 4CL1 expression，promotes lignification	枇杷 Eriobotrya japonica	Zhang et al.,2018
	低温 Cold	—	MYB5/12/13/ 14	激活C4H1/4CL3，促进木质化 Activates C4H1/4CL3，promotes lignification	猕猴桃 Actinidia chinensis	焦健青,2022
	低温 Cold	—	MYB8	激活4CL1，促进木质化 Activates 4CL1 expression，promotes lignification	枇杷 Eriobotrya japonica	Wang et al.,2016
	低温 Cold	—	MYB15	激活4CL2，导致木质素积累，引发汁胞粒化 Activates 4CL2 expression，leads to lignin accumulation，triggers juice sac granulation	甜橙 Citrus sinensis	Song et al.,2023
	热激 Heat	—	MYB63	激活PME3/PG1，加速细胞壁降解，加重冷害 Activates PME3/PG1，accelerates cell wall degradation，aggravates chilling injury	西葫芦 Cucurbita pepo	Zuo et al.,2025
	低温 Cold	—	NAC1	激活PAL1/4CL1，促进木质化 Activates PAL1/4CL1，promotes lignification	枇杷 Eriobotrya japonica	Xu et al.,2015
	低温 Cold	—	NAC1	激活CESA7/6B，促进细胞壁增厚，增强抗冷性 Activates CESA7/6B，promotes cell wall thickening，enhances chilling tolerance	香蕉 Musa acuminata	Yin et al.,2024
	低温 Cold	—	NAC3	激活CAD-like，促进木质素合成 Activates CAD-like，promotes lignin synthesis	枇杷 Eriobotrya japonica	Ge et al.,2017
	低温 Cold	—	NAC76	激活CAD1，增强细胞壁强度与抗冷能力 Activates CAD1，enhances cell wall strength and chilling tolerance	辣椒 Capsicum annuum	Xiao et al.,2025
	PSK-α	bHLH77	WRKY33	激活PAL/C4H/4CL/POD，促进木质化 Activates PAL/C4H/4CL/POD，promotes lignification	枇杷 Eriobotrya japonica	Liu et al.,2026
	低温 Cold	—	WRKY65	抑制4CL7/CAD1/COMT/F5H，减轻木质化 Suppresses 4CL7/CAD1/COMT/F5H expression，alleviates lignification	桃 Prunus persica	Wang et al.,2024