Advances in Chilling Injury and Associated Quality Deterioration in Postharvest Fruits

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

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 1871-1890.doi: 10.16420/j.issn.0513-353x.2026-0388

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Advances in Chilling Injury and Associated Quality Deterioration in Postharvest Fruits

LIU Mengfei, LI Junduo, HU Ruoqian, SHI Yanna, XU Changjie^*(), CHEN Kunsong

College of Agriculture & Biotechnology， Zhejiang University; Zhejiang Provincial Key Laboratory of Horticultural Crop Quality Manipulation;Ministry of Agriculture and Rural Affairs Laboratory of Horticultural Plant Crop Growth and Development， Hangzhou 310058， China

Received:2026-05-08 Revised:2026-06-12 Online:2026-07-23 Published:2026-07-23
Contact: XU Changjie

Abstract

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

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.

Figures/Tables 4

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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

Advances in Chilling Injury and Associated Quality Deterioration in Postharvest Fruits

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