Recent Advances in JAs-Induced Improvement of Postharvest Chilling Resistance in Horticultural Crops

doi:10.16420/j.issn.0513-353x.2024-0969

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (8): 1987-2020.doi: 10.16420/j.issn.0513-353x.2024-0969

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Recent Advances in JAs-Induced Improvement of Postharvest Chilling Resistance in Horticultural Crops

TAN Xiaoyu¹, NIU Junpeng²^,³, WANG Quanzhi¹, ZHANG Li², ZHU Lijuan², WANG Guodong³, ZHENG Heyun⁴, YU Zhifang², DUAN Yuquan⁵, JIANG Li², SUN Xiuxiu⁶, YANG Ying⁷, LUO Weiqi⁶, LI Xuehui², GUAN Le², ZHAO Yanling¹, LI Guoxiao¹, YIN Congfei¹, GE Cheng¹, MA Min², JIA Luting², ZHANG Xu², ZHAO Yaoyao⁵^,^*(), GENG Xinli⁴^,^*(), WANG Libin²^,⁸^,^*(), and ZHANG Shaoling²^,^*()

¹ School of Agronomy and Horticulture， Jiangsu Vocational College of Agriculture and Forestry，Zhenjiang， Jiangsu 212499， China
² Nanjing Agricultural University， Nanjing 210095， China
³ College of Life Science， Shaanxi Normal University， Xi’an 710119， China
⁴ Research Institute of Grape and Melon of Xinjiang Uygur Autonomous， Turpan， Xinjiang 838200， China
⁵ Institute of Food Science and Technology， Chinese Academy of Agricultural Sciences， Beijing 100193
⁶ U.S. Horticultural Research Laboratory， Ft. Pierce， Florida 34945， USA
⁷ College of Horticulture， Anhui Agricultural University， Hefei 230036， China
⁸ School of Food and Biological Engineering， Hezhou University，Hezhou， Guangxi 542899， China

Received:2025-06-13 Revised:2025-07-15 Online:2025-08-25 Published:2025-08-19
Contact: ZHAO Yaoyao, GENG Xinli, WANG Libin, and ZHANG Shaoling

Abstract

Abstract:

Low-temperature（0-15 ℃）storage would trigger a series of the complicated physio- biochemical reactions in horticulture crops，damage the integrity，fluidity，and functionality （semi-permeability）of cell membranes，thereby causing chilling injury. To mitigate chilling damage，the CBF-(in)dependent signal transduction pathways are initiated in plants；among them，the CE1-CBF-COR transcriptional cascade is of great importance in the CBF-dependent signal transduction pathway. In addition，some endogenous hormones such as jasmonic acid（JA），ethylene（ETH）and abscisic acid（ABA） also rapidly accumulate during plant response to low temperature. JA and its derivatives are collectively referred to as JAs，which are formed after oxidation，cyclization，reduction，β-oxidation，modification and activation of α-linolenic acid or hexadecatrienoic acid. Upon JAs synthesis followed by perception by COI1，the transcription factor MYC2 is released from the repressor protein JAZs，which was ubiquitinated for degradation，to activate the downstream signal transduction process，causing the induction of several physio-biochemical alternations（including increment of antioxidant biosynthesis，improvement of antioxidant enzyme activity，inhibition of lipid metabolism，enhancement of arginine metabolism，promotion of energy metabolism，and activation of other plant hormone signal transduction pathways）. The abovementioned responses would help to maintain the integrity，fluidity，and functionality（semi-permeability）of cell membrane，and thus improve the capacity of horticultural crops postharvest to resist low temperature stress. In recent years，studies have revealed that：（a）MYC2 regulates the transcription of downstream target genes by forming homodimers or heterodimers with MYC3 and MYC4；（b）BBX37 can activate the transcription of downstream CBFs genes independently or in synergy with ICE1；（c）MED16 enhances the stability of MED25，positively regulating the transcriptional output of JAs signaling，among other functions. This paper first elaborates on the characteristics of postharvest chilling injury in horticultural crops，reviews the roles of CBF-(in)dependent signal transduction pathways in plant responses to low-temperature stress，summarizes the biosynthesis，signaling pathways，and regulatory mechanisms of JAs，and introduces the applications and mechanisms of action of JAs in alleviating chilling injury in horticultural crops. Based on these，this paper systematically summarizes the association between the JAs signaling pathway and CBF-(in)dependent signal transduction pathways during low-temperature stress and proposes future research directions. Future research directions should focus on：（a）delving deeper into and validating the associations between JAs signaling pathways and CBF-(in)dependent signaling cascades；（b）elucidating the roles of post-transcriptional processing of genes and post-translational modifications of proteins in the synthesis or signaling transduction of JAs；（c）exploring the synergistic effects of JAs with other endogenous plant hormones in resistance to low temperature stress；and（d）dissecting the mechanisms of newly discovered components（MYC3/4，MED16，BBX37，etc.）in the low temperature response process.

Key words: postharvest, low-temperature stress, CBF-(in)dependent signaling pathways, JAs biosynthetic and transduction pathways, regulatory mechanisms, relationship

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, and ZHANG Shaoling. Recent Advances in JAs-Induced Improvement of Postharvest Chilling Resistance in Horticultural Crops[J]. Acta Horticulturae Sinica, 2025, 52(8): 1987-2020.

Figures/Tables 7

Table 1 Chilling injury symptoms of different fruits and vegetables with their critical chilling temperatures

园艺作物 Horticultural crop	冷害症状 Chilling injury symptoms	临界冻害温度/℃ Critical chilling temperature
果树Fruit tree
苹果（亚热带季风气候区） Apple（Subtropical monsoon climate region）	内部变褐，褐芯，脆化，软烫伤 Internal browning，brown core，soggy breakdown，soft scald	2 ~ 3
凤梨释迦Atemoya	表皮变黑，不成熟，果肉变色Skin darkening，failure to ripen，pulp discoloration	4
油梨Avocado	果肉灰褐色褪色Grayish-brown discoloration of flesh	4.5 ~ 13
木橘Bael	表皮出现褐色斑点Brown spots on skin	3
香蕉（绿色或成熟） Banana（green or ripe）	成熟时颜色暗淡Dull color when ripened	11.5 ~ 13
面包果Breadfruit	非正常成熟，暗褐色褪色Abnormal ripening，dull brown discoloration	7 ~ 12
蔓越莓Cranberry	橡胶质地，红褐色果肉Rubbery texture，red flesh	2
番石榴Guava fruit	果肉损伤，腐烂Pulp injury，decay	4.5
西柚Grapefruit	烫伤痕迹，点状腐蚀，水渍Scald，pitting，watery breakdown	10
柠檬Lemon	点状腐蚀，膜质染色，红色斑点Pitting，membranous staining，red blotch	11 ~ 13
酸橙Lime	点状腐蚀，随着时间推移变成棕褐色Pitting，turning tan with time	7 ~ 9
荔枝Lychee	表皮变褐Skin browning	3
杧果Mango	表皮呈灰鳞片状褪色，成熟不均匀 Grayish scald-like discoloration of skin，uneven ripening	10 ~ 13
山竹Mangosteen	皮层硬化褐变Hardening and browning of the cortex	4 ~ 8
新鲜橄榄Olive，fresh	内部变褐Internal browning	7
橘Orange	点状腐蚀，棕色污渍Pitting，brown stain	3
木瓜Papaya	不成熟，变味，腐烂Pitting，failure to ripen，off-flavor，decay	7
西番莲Passion fruit	表皮褪色成暗红色，味道变淡，腐烂 Dark red discoloration on skin，loss of flavor，decay	10
菠萝Pineapple	成熟后呈暗绿色Dull green when ripened	7 ~ 10
石榴Pomegranate	点状腐烂，内外部褐变Pitting，external and internal browning	4.5
红毛丹Rambutan	外果皮变黑Darkening of exocarp	10
树番茄Tamarillo	表面点状腐蚀，褪色Surface pitting，discoloration	3 ~ 4
瓜类Melons and gourds
哈密瓜Cantaloup	点状腐蚀，表面腐化Pitting，surface decay	2 ~ 5
网纹瓜Crenshaw and Persian	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
蜜瓜Honeydew	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
白兰瓜Casaba	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
西瓜Watermelon	点状腐蚀，异味Pitting，objectionable flavor	4.5
蔬菜（果菜）Vegetables（fruits and vegetables）
佛手瓜Choyote	暗褐色褪色，点状腐蚀，果肉变黑Dull brown discoloration，pitting，flesh darkening	5 ~ 10
黄瓜Cucumber	点状腐蚀，水渍状斑点，腐烂Pitting，water-soaked spots，decay	7
茄子Eggplant	表皮有烫伤痕迹，黑斑病，种子变黑Surface scald，alternaria rot，blackening of seeds	7
豆薯Jicama	表面腐烂，褪色Surface decay，discoloration	13 ~ 18
秋葵Okra pod	褪色，水浸区域，点状腐蚀，腐烂Discoloration，water-soaked areas，pitting，decay	7
南瓜和硬壳南瓜 Pumpkin and hard-shell squash	腐烂，特别是黑斑病Decay，especially alternaria rot	10
利马豆Phaseolus lunatus	锈褐色斑块、斑点或区域Rusty brown specks，spots，or areas	0 ~ 2
菜豆Phaseolus vulgaris	点状腐蚀和褐斑Pitting and russeting	7
辣椒（甜）Pepper（sweet）	豆荚和花萼上的片状腐烂，黑斑病，种子变黑 Sheet pitting，alternaria rot on pods and calyxes，darkening of seed	7
西红柿（成熟期）Tomato（ripe）	浸水，软化，腐烂Water-soaking and softening，decay	7 ~ 10
西红柿（绿熟期）Tomato（mature green）	成熟时色泽差，黑斑病Poor color when ripe，alternaria rot	13
蔬菜（叶菜）Vegetables（leafy vegetables）
石刁柏Asparagus	暗淡，灰绿色，尖端软弱无力Dull，gray-green，and limp tips	0 ~ 2
蕹菜Water convolvulus	茎叶变黑Darkening of leaves and stems	10

Table 1 Chilling injury symptoms of different fruits and vegetables with their critical chilling temperatures

园艺作物 Horticultural crop	冷害症状 Chilling injury symptoms	临界冻害温度/℃ Critical chilling temperature
果树Fruit tree
苹果（亚热带季风气候区） Apple（Subtropical monsoon climate region）	内部变褐，褐芯，脆化，软烫伤 Internal browning，brown core，soggy breakdown，soft scald	2 ~ 3
凤梨释迦Atemoya	表皮变黑，不成熟，果肉变色Skin darkening，failure to ripen，pulp discoloration	4
油梨Avocado	果肉灰褐色褪色Grayish-brown discoloration of flesh	4.5 ~ 13
木橘Bael	表皮出现褐色斑点Brown spots on skin	3
香蕉（绿色或成熟） Banana（green or ripe）	成熟时颜色暗淡Dull color when ripened	11.5 ~ 13
面包果Breadfruit	非正常成熟，暗褐色褪色Abnormal ripening，dull brown discoloration	7 ~ 12
蔓越莓Cranberry	橡胶质地，红褐色果肉Rubbery texture，red flesh	2
番石榴Guava fruit	果肉损伤，腐烂Pulp injury，decay	4.5
西柚Grapefruit	烫伤痕迹，点状腐蚀，水渍Scald，pitting，watery breakdown	10
柠檬Lemon	点状腐蚀，膜质染色，红色斑点Pitting，membranous staining，red blotch	11 ~ 13
酸橙Lime	点状腐蚀，随着时间推移变成棕褐色Pitting，turning tan with time	7 ~ 9
荔枝Lychee	表皮变褐Skin browning	3
杧果Mango	表皮呈灰鳞片状褪色，成熟不均匀 Grayish scald-like discoloration of skin，uneven ripening	10 ~ 13
山竹Mangosteen	皮层硬化褐变Hardening and browning of the cortex	4 ~ 8
新鲜橄榄Olive，fresh	内部变褐Internal browning	7
橘Orange	点状腐蚀，棕色污渍Pitting，brown stain	3
木瓜Papaya	不成熟，变味，腐烂Pitting，failure to ripen，off-flavor，decay	7
西番莲Passion fruit	表皮褪色成暗红色，味道变淡，腐烂 Dark red discoloration on skin，loss of flavor，decay	10
菠萝Pineapple	成熟后呈暗绿色Dull green when ripened	7 ~ 10
石榴Pomegranate	点状腐烂，内外部褐变Pitting，external and internal browning	4.5
红毛丹Rambutan	外果皮变黑Darkening of exocarp	10
树番茄Tamarillo	表面点状腐蚀，褪色Surface pitting，discoloration	3 ~ 4
瓜类Melons and gourds
哈密瓜Cantaloup	点状腐蚀，表面腐化Pitting，surface decay	2 ~ 5
网纹瓜Crenshaw and Persian	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
蜜瓜Honeydew	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
白兰瓜Casaba	红褐色褪色，麻点，表面腐烂，不成熟 Reddish-tan discoloration，pitting，surface decay，failure to ripen	7 ~ 10
西瓜Watermelon	点状腐蚀，异味Pitting，objectionable flavor	4.5
蔬菜（果菜）Vegetables（fruits and vegetables）
佛手瓜Choyote	暗褐色褪色，点状腐蚀，果肉变黑Dull brown discoloration，pitting，flesh darkening	5 ~ 10
黄瓜Cucumber	点状腐蚀，水渍状斑点，腐烂Pitting，water-soaked spots，decay	7
茄子Eggplant	表皮有烫伤痕迹，黑斑病，种子变黑Surface scald，alternaria rot，blackening of seeds	7
豆薯Jicama	表面腐烂，褪色Surface decay，discoloration	13 ~ 18
秋葵Okra pod	褪色，水浸区域，点状腐蚀，腐烂Discoloration，water-soaked areas，pitting，decay	7
南瓜和硬壳南瓜 Pumpkin and hard-shell squash	腐烂，特别是黑斑病Decay，especially alternaria rot	10
利马豆Phaseolus lunatus	锈褐色斑块、斑点或区域Rusty brown specks，spots，or areas	0 ~ 2
菜豆Phaseolus vulgaris	点状腐蚀和褐斑Pitting and russeting	7
辣椒（甜）Pepper（sweet）	豆荚和花萼上的片状腐烂，黑斑病，种子变黑 Sheet pitting，alternaria rot on pods and calyxes，darkening of seed	7
西红柿（成熟期）Tomato（ripe）	浸水，软化，腐烂Water-soaking and softening，decay	7 ~ 10
西红柿（绿熟期）Tomato（mature green）	成熟时色泽差，黑斑病Poor color when ripe，alternaria rot	13
蔬菜（叶菜）Vegetables（leafy vegetables）
石刁柏Asparagus	暗淡，灰绿色，尖端软弱无力Dull，gray-green，and limp tips	0 ~ 2
蕹菜Water convolvulus	茎叶变黑Darkening of leaves and stems	10

Fig. 1 Physio-biochemical responses of horticultural crops to chilling temperature Wang，1989；Paull，1990；Theocharis et al.，2012. Cryogenic temperature triggers a series of the complicated physio-biochemical alternations in plant cell，leading to the occurrence of chilling injury symptom. Firstly，cell membrane was changed from liquid crystal phase to gelatinous phase，resulting in the loss of membrane semi-permeability，disfunction of membrane proteins，decrement of protoplasm fluidity，and so on. In accompany with the occurrence of electrolyte leakage，energy reduction，and oxidative stress，cell membrane damage is exacerbated. If the chilling-exposure continues，cell membrane would be irreversibly damaged，causing chilling injury symptom

Fig. 2 CBF-dependent signaling pathway in plants Li et al.，2017a；Liu et al.，2017b；Ding et al.，2019. The ICE1-CBF-COR transcriptional cascade is of great importance for the CBF-dependent signaling pathway. During chilling stress，the membrane protein complex COLD1-RGA1 senses environmental signals to induce Ca2+ influx. After perception of Ca2+ signals，CRLK and EGR2-OST1-ICE1 signaling pathways are activated to enhance the stability and transcriptional activity of ICE1，resulting in the upregulated expression levels of CBFs. On the other hand，the CRPK1 signal transduction pathway，which negatively regulates the stability of CBFs，is also initiated to avoid the cellular over-response to low temperature. After translation，CBFs could bind to cis-acting element（CRT/DRE）in the promoters of the downstream CORs genes and then initiate their transcription. CBF：C-repeat binding factors；COLD1：Chilling tolerance divergence 1；COR：Cold responsive gene；CRLK：Calcium/calmodulin-regulated receptor-like kinase；CRPK1：Cold-responsive protein kinase 1；EGR2：Clade-E growth-regulating 2；MAP3K：Mitogen-activated protein kinase kinase kinase；MEKK：MAPK/ERK Kinase Kinase；MKK：MAPK Kinase；MPK：Mitogen-activated protein kinase；ICE1：Inducer of CBF expression 1；OST1：Open stomata 1； P：Phosphorylation；RGA1：Rice G-protein α subunit 1

Fig. 3 Biosynthetic pathway of JAs in plants Sharma & Laxmi，2016；Ali & Baek，2020；Wan & Xin，2022. JAs is derived from α-LeA and HTA via oxidation，cyclization，reduction，β-oxidation，modification and activation. Upon release from chloroplast membrane，α-LeA and HTA are catalyzed by LOX，AOS and AOC into 12-OPDA and dn-OPDA，which are then transported into peroxisome by ABC transporter or ionic barrier. In peroxisome，12-OPDA and dn-OPDA are converted into (+)-7-iso-JA via the actions of several enzymes，including OPR2/3，OPCL1，ACX，MFP and KAT. After transportation into the cytoplasm，(+)-7-iso-JA could be catalyzed by JAR1 and JMT to form JA-Ile and MeJA，respectively. ACX：Acyl-CoA oxidase；AOC：Alle-neoxide cyclase；AOS：Allene oxide synthase；4,5-didehydro-JA：4,5-Didehydro-jasmonic acid；dn-OPDA：Dinor OPDA；10,11-EHT：10,11(S)-epoxy- hexadeca(tri)enoic acid；12,13-EOT：12,13-(S)-epoxy-octadecatrienoic acid；11-HPHT：11-Hydroperoxyhexadecatrienoic acid；13-HPOT：13-Hydroperoxyoctadecatrienoic acid；HTA：Hexadecatrienoic acid；(+)-7-iso-JA：(+)-7-iso-Jasmonic acid；JA-Ile：Jasmonoyl-isoleucine；JAR1：Jasomonate resistant 1；JMT：Jasmonic acid carboxyl methyltransferase；KAT：3-Ketoacyl-CoA thiolase；α-LeA：α-Linolenic acid；LOX：Lipoxygenases；OPC-8：3-Oxo-2-((Z)-pent-2-en-1-yl)cyclopentane-1-octanoic acid；MeJA：Methyl jasmonate；MFP：Multifunctional protein；OPC：8-CoA：3-Oxo-2-((Z)-pent-2-en-1-yl)cyclopentane-1-octanoyl-CoA；OPCL1：OPC-8 coenzyme A ligase1；12-OPDA：12-Oxo-phytodienoic acid；OPR：OPDA reductase；PLs：Phospholipases；tn-OPDA：Tetranor-OPDA

Fig. 4 JAs signal transduction process in plants Sharma & Laxmi，2016；Ali & Baek，2020；Li et al.，2022. JA-Ile is transported from cytosol into the nucleus with the aid of JAT1 to initiate its signal transduction process. When JA-Ile is low，the repressor protein JAZs recruits NINJA，TPL，and HDA6/19 to form a closed transcriptional inhibition complex to inhibit MYC2 transcriptional activity. At the specific developmental stages or under（a）biotic stress，the rapid accumulation of JA-Ile in the nucleus promotes the interaction between JAZs and COI1 in the E3 ubiquitin ligase complex SCFCOI1，leading to the ubiquitin-mediated degradation of JAZs by IP5 and 26S proteasome. Then，MYC2 is released from JAZs. Then，MYC2 forms homodimers or heterodimers with its homologs MYC3 and MYC4，which bind to the G-box element. Subsequently，MED25 recruits General Transcription Factors（GTFs）and RNA polymerase II to the promoter to initiate transcription. Additionally，MED16 enhances the stability of MED25 through interaction with it，which is crucial for activating the expression of JAs-responsive genes. COI1：Coronatine insensitive 1；CUL：Cullin；E1，E2：Ubiquitin-conjugating enzymes；GTF：General transcription factor；HDA6，HDA19：Histone deacetylase 6，19；JA：Jasmonic acid；JA-Ile：Jasmonyl isoleucine；JAT1：Jasmonic acid transfer protein 1；JAZ：Jasmonate ZIM domain；MED16，25：Mediator complex subunit 16；MYC：Myelocytomatosis transcription factor；NINJA：Novel interactor of JAZ；RBX1：Ring box 1；RNA Pol II：RNA polymerase II；SCF：Skp1-cullin-F-box e3 ubiquitin ligase complex；SKP1：S-phase kinase-associated protein 1；TPL：Topless；Ubi：Ubiquitin

Fig. 5 Synthesis and metabolic pathways of some substances in plants a：The biosynthetic pathways of AsA and GSH in plant（Forman et al.，2009；Ishikawa et al.，2018）.（I）D-mannose/L-galactose pathway（also known as Smirnoff Wheeler pathway）is the most important pathway for AsA biosynthesis in plant. D-glucose-6-phosphate in cytosol is converted into L-galactono-1,4-lactone by GPI，PMI，PMM，GMP，GME，GDP-GGP，GPP，and GalDH；after transportation into mitochondria，L-galactono-1,4-lactone is oxidized into AsA with the aid of GalLDH.（II）GSH is derived from glutamic acid through the actions of glutamate-cysteine ligase and glutathione synthase. b：The biosynthetic pathways of carotenoids and abscisic acid in plant（Nurbekova et al.，2021；Sun et al.，2022；Wu et al.，2022b）. The biosynthesis of carotenoids and abscisic acid in plant begins with MEP pathway. Geraniol pyrophosphate，which is produced via MEP pathway，is catalyzed by PSY，PDS，ZISO，ZDS and CRTISO to produce lycopene. The latter is then cyclized by LCYE and LCYB to form β-carotene and α-carotene. β-carotene could further be converted into zeaxanthin，violaxanthin，neoxanthin，9-cis-violaxanthin and 9’-cis-neoxanthin with the aid of various enzymes，including BCH/CrtZ，ZEP，NSY and uncharacterized isomerase. Moreover，9-cis-violaxanthin and 9’-cis-neoxanthine，which are derived from the carotenoid-biosynthetic pathway，could be acted as the substrates for ABA formation through the catalytic actions of NCED，ABA2 and AAO3. c：Reactive oxygen species scavenging pathways in plant（Hasanuzzaman et al.，2022）.（Ⅰ）Ascorbate-glutathion cycle；（Ⅱ）superoxide dismutase-glutathion peroxidase cycle；（Ⅲ）the catalase pathway. d：Arginine metabolic pathway in plant（Cao et al.，2012；Zhang et al.，2012；Winter et al.，2015）. Putrescine in plant is produced from arginine via ADC-AIH-NLP and ARG-ODC pathways. After formation，putrescine would further be converted by SPDS and SPMS into spermidine and spermidine. Ornithine，as an metabolic intermediate during putrescine formation，could be acted as the substrate for arginine regeneration，with the aid of OTC，ASSY and ASL，to maintain its level in plant cell；additionally，ornithine might also be catalyzed by δOAT，P5CS，GAD，P5CDH and P5CR，producing γ-aminobutyric acid or proline. AAO3：Abscisic-aldehyde oxidase；ABA2：Xanthoxin dehydrogenase；ADC：Arginine decarboxylase；ADP：Adenosine diphosphate；AIH：Agmatine iminohydrolase；APX：Ascorbate peroxidase；ARG：Arginase；AsA：Ascorbic acid；ASL：Argininosuccinate lyase；ASSY：Argininosuccinate synthase；ATP：Adenosine triphosphate；BCH/CrtZ：β-Carotenehydroxylase；CRTISO：Carotene isomerase；DHA：Dehydroascorbate；DHAR：Dehydroascorbate reductase；GAD：Glutamate decarboxylase；GalDH：L -galactose dehydrogenase；GalLDH：L -galactono-1,4-lactone dehydrogenase；GCL：Glutamate cysteine ligase；GGP：GDP- L -galactose phosphorylase；GME：GDP-mannose-3’，5’-epimerase；GMP：GDP-mannose pyrophosphorylase；GPI：Glucose-6-phosphate isomerase；GPP：L-galactose-1-phosphate phosphatase；GPX： Glutathione peroxidase；GR：Glutathione reductase；GSA：Glutamic-y-semialdehyde；GSH：Glutathione；GSS：Glutathione synthase；GSSG：oxidized glutathione；H2O2：Hydrogen peroxide；LCYB：Lycopeneβ-cyclase；LCYE：Lycopene ε-cyclase；MDHAR：Monodehydroascorbate reductase；MEP：Methylerythritol phosphate；NCED：9-cis-epoxycarotenoid dioxygenase；NLP：N-carbamoylputrescine amidase；NOS：NO synthase；NSY：Neoxanthinsynthase；δOAT：Ornithine-δ-aminotransferase；ODC：Ornithine decarboxylase；OTC：Ornithine transcarbamylase；PDS：Phytoene desaturase；Pi：Inorganic phosphate；PMI：Mannose-6-phosphate isomerase；PPM：Phosphomanno mutase；ProDH：Proline dehydrogenase；PSY：Phytoene synthase；P5C：Pyrroline-5-carboxylate；P5CDH：Pyrroline-5-carboxylate dehydrogenase；P5CR：Pyrroline-5-carboxylate reductase；P5CS：Pyrroline-5-carboxylate synthetase；SOD：Superoxide dismutase；SPDS：Spermidine synthase；SPMS：Spermine synthase；ZDS：ζ-Carotene desaturase；ZEP：Zeaxanthinepoxidase；ZISO：ζ-Carotene isomera

Fig. 6 Mechanism on the JAs-induced improvement of chilling resistance in horticultural crops Bolt et al.，2017；Sun et al.，2019；An et al.，2021，2022；Song et al.，2022. Low temperature induces JAs biosynthesis. Afterwards，JAs could directly or indirectly activate the CBF-(in)dependent signal transduction pathway with the aid of several signaling transduction elements，triggering a series of the complicated physio-biochemical alternation（including increment of antioxidant biosynthesis，improvement of antioxidant enzyme activity，inhibition of lipid metabolism，enhancement of arginine metabolism，promotion of energy metabolism，and activation of other plant hormone signal transduction pathways）. The abovementioned responses could maintain the integrity，fluidity，and functionality（semi-permeability）of cell membrane，thereby improving the resistance of horticultural crops to chilling stress. ABA：Abscisic acid；ABI4：ABA insensitive 4；ABI5-like：ABA insensitive 5-like；ACX：Acyl-CoA oxidase；AOC：Alle-neoxide cyclase；AOS：Allene oxide synthase；BBX37：B-box protein BBX37；CBF：C-repeat binding factors；COI1：Coronatine insensitive 1；COR：Cold responsive genes；CUL：Cullin；4,5-didehydro-JA：4,5-Didehydro- jasmonic acid；dn-OPDA：Dinor OPDA；E1，E2：Ubiquitin-conjugating enzymes；10,11-EHT：10，11(S)-epoxy-hexadeca (tri)enoic acid；12,13-EOT：12,13-(S)-epoxy-octadecatrienoic acid；ERF：Ethylene responsive factor；ETH：Ethylene；11-HPHT：11-Hydroperoxyhexadecatrienoic acid；13-HPOT：13-Hydroperoxyoctadecatrienoic acid；HTA：Hexadecatrienoic acid；ICE1：Inducer of CBF expression 1；(+)-7-iso-JA：(+)-7-iso-Jasmonic acid；JA-Ile：Jasmonoyl-isoleucine；JAR1：Jasomonate resistant 1；JAT1：Jasmonic acid transfer protein 1；JAZ：Jasmonate ZIM domain；JMT：Jasmonic acid carboxyl methyltransferase；KAT：3-Ketoacyl-CoA thiolase；α-LeA：α-Linolenic acid；LOX：Lipoxygenases；MeJA：Methyl jasmonate；MFP：Multifunctional protein；MYC：Myelocytomatosis transcription factor；NINJA：Novel interactor of JAZ；OPC-8：3-Oxo-2-((Z)-pent-2-en-1-yl）cyclopentane-1-octanoic acid；OPC：8-CoA：3-Oxo-2-（(Z)-pent-2-en-1-yl）cyclopentane-1-octanoyl-CoA；OPCL1：OPC-8 coenzyme A ligase1；12-OPDA：12-Oxo-phytodienoic acid；OPR：OPDA reductase；PLs：Phospholipases；RBX1：Ring box 1；RNA Pol II：RNA polymerase II；SKP1：S-phase kinase-associated protein 1；tn-OPDA：Tetranor-OPDA；TPL：Topless；Ubi：Ubiquitin

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Recent Advances in JAs-Induced Improvement of Postharvest Chilling Resistance in Horticultural Crops

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