褪黑素对冷藏后芒果果实冷害和成熟的影响及生理机制

摘要/Abstract

摘要： 以采后‘贵妃’芒果为试材，在0.5 mmol/L褪黑素（MT）溶液中浸泡1 h，晾干后置于4 °C下贮藏28 d，随后转移至室温（25 °C）贮藏4 d，调查MT对冷藏后芒果果实冷害（CI）和成熟的影响及相关机理。结果显示，MT处理有效降低了芒果果实的CI指数，提高了可溶性固形物含量、呼吸速率和乙烯释放量并加速了色相值（h°）和硬度下降；MT处理促进了水溶性果胶和螯合性果胶含量升高，提高了多聚半乳糖醛酸酶、β-半乳糖苷酶和纤维素酶活性，加速了果胶甲酯酶活性下降，从而促进了细胞壁降解。此外，MT处理提高了果实1-氨基环丙烷-1-羧酸（ACC）合成酶（ACS）和ACC氧化酶（ACO）活性及其编码基因MiACS和MiACO的表达水平，同时上调了MiETR1、MiERS1、MiERF1、MiEIN2、MiCBF1和MiICE1并下调MiCTR1等基因的表达。上述结果表明MT能有效改善冷藏后芒果后熟恢复能力并提高果实抗冷性。

关键词: 芒果, 褪黑素, 冷害, 货架期, 细胞壁代谢, 乙烯生物合成

Abstract: Using postharvest ‘Guifei’ mango fruit as experimental material, we investigated the effects of 0.5 mmol/L melatonin (MT) treatment on chilling injury (CI) and ripening and related mechanism in fruit during shelf life at 25 °C after refrigeration at 4 °C. The results showed that MT treatment markedly decreased CI index, enhanced soluble solids content (SSC), respiration rate and ethylene production, while accelerating the declines in hue angle (h°) and firmness in chilled mango fruit. Compared to the control fruit, MT treatment promoted the increases in contents of water-soluble pectin and chelate (CDTA)-soluble pectin during shelf life in mango fruit after refrigeration. MT treatment promoted the increases in activities of polygalacturonase (PG), β-galactosidase (β-Gal) and endo-1,4-β-D-glucanase (EGase) and accelerated the decrease in pectin methylesterase (PME) activity, thereby leading to cell wall degradation. Compared to control fruit, higher activities of 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase (ACS) and ACC oxidase (ACO) and the expressions of their encoding genes (MiACS and MiACO) were observed in MT-treated fruit. Moreover, MT up-regulated the expression of MiETR1, MiERS1, MiERF1, MiEIN2, MiCBF1 and MiICE1 while down-regulating the expression of MiCTR1. These results indicate that MT treatment could promote ripening recovery and improve the chilling tolerance in mango fruit during shelf life after removal of low temperature.

Key words: mango, melatonin, chilling injury, shelf life, cell wall metabolism, ethylene biosynthesis

中图分类号:

TS255.36

徐萍黄婷刘士琦胡美姣高兆银刘家粮张正科. 褪黑素对冷藏后芒果果实冷害和成熟的影响及生理机制[J]. 食品科学.

萍徐 jialiang LIU Zhengke .ZHANG. Effect of Melatonin on Chilling Injury and Ripening in Postharvest Mango Fruit During Shelf Life after Refrigeration[J]. FOOD SCIENCE.

参考文献

[1] SIVAKUMAR D, JIANG Yueming, YAHIA E M. Maintaining mango (Mangifera indica L.) fruit quality during the export chain[J]. Food Research International, 2011, 44(5): 1254-1263. DOI:10.1016/j.foodres.2010.11.022.
[2] LE T D, NGUYEN T V, MUOI N V, et al. Supply chain management of mango (Mangifera indica L.) fruit: A review with a focus on product quality during postharvest[J]. Frontiers in Sustainable Food Systems, 2022, 5: 571. DOI:10.3389/fsufs.2021.799431.
[3] RAO D V S, SHIVASHANKARA K S. Individual shrink wrapping extends the storage life and maintains the antioxidants of mango (cvs. ‘Alphonso’ and ‘Banganapalli’) stored at 8 °C[J]. Journal of Food Science and Technology, 2015, 52: 4351-4359. DOI:10.1007/s13197-014-1468-6.
[4] ZHANG Zhengke, GAO Zhaoyin, LI Min, et al. Hot water treatment maintains normal ripening and cell wall metabolism in mango (Mangifera indica L.) fruit[J]. HortScience, 2012, 47(10): 1466-1471. DOI:10.21273/HORTSCI.47.10.1466.
[5] ZHANG Zhengke, ZHU Qinggang, HU Meijiao, et al. Low-temperature conditioning induces chilling tolerance in stored mango fruit[J]. Food Chemistry, 2017, 219: 76-84. DOI:10.1016/j.foodchem.2016.09.123.
[6] JIANG Long, HAN Zhanhong, LIU Jialiang, et al. Intermittent stepwise cooling and warming ameliorate chilling injury and improve quality in postharvest ‘Guifei’ mango fruit[J]. LWT, 2023, 181: 114740. DOI:10.1016/j.lwt.2023.114740.
[7] 巩彪, 史庆华. 园艺作物褪黑素的研究进展[J]. 中国农业科学, 2017, 50(12): 12. DOI:10.3864/j.issn.0578-1752.2017.12.013.
[8] ARNAO M B, HERNA?NDEZ-RUIZ J. Melatonin and reactive oxygen and nitrogen species: a model for the plant redox network[J]. Melatonin Research, 2019, 2(3): 152-168. DOI:10.32794/mr11250036.
[9] ARNAO M B, CANO A, HERNáNDEZ-RUIZ J. Phytomelatonin: an unexpected molecule with amazing performances in plants[J]. Journal of Experimental Botany, 2022, 73(17): 5779-5800. DOI:10.1093/jxb/erac009.
[10] SHAH H M S, SINGH Z, AFRIFA-YAMOAH E, et al. Insight into the role of melatonin in mitigating chilling injury and maintaining the quality of cold-stored fruits and vegetables[J]. Food Reviews International, 2023: 1-27. DOI:10.1080/87559129.2023.2212042.
[11] BHARDWAJ R, AGHDAM M S, ARNAO M B, et al. Melatonin alleviates chilling injury symptom development in mango fruit by maintaining intracellular energy and cell wall and membrane stability[J]. Frontiers in Nutrition, 2022, 9. DOI:10.3389/fnut.2022.936932.
[12] BHARDWAJ R, PAREEK S, DOMíNGUEZ-AVILA J A, et al. An exogenous pre-storage melatonin alleviates chilling injury in some mango fruit cultivars, by acting on the enzymatic and non-enzymatic antioxidant system[J]. Antioxidants, 2022, 11(2): 384. DOI:10.3390/antiox11020384.
[13] BHARDWAJ R, PAREEK S, SARAVANAN C, et al. Contribution of pre-storage melatonin application to chilling tolerance of some mango fruit cultivars and relationship with polyamines metabolism and γ-aminobutyric acid shunt pathway[J]. Environmental and Experimental Botany, 2022, 194: 104691. DOI:10.1016/j.envexpbot.2021.104691.
[14] XU Ping, HUBER D J, GONG Di, et al. Amelioration of chilling injury in ‘Guifei’ mango fruit by melatonin is associated with regulation of lipid metabolic enzymes and remodeling of lipidome[J]. Postharvest Biology and Technology, 2023, 198: 112233. DOI:10.1016/j.postharvbio.2022.112233.
[15] HU Meijiao, YANG Dongping, HUBER D J, et al.Reduction of postharvest anthracnose and enhancement of disease resistance in ripening mango fruit by nitric oxide treatment[J].Postharvest Biology and Technology, 2014, 97(11): 115-122. DOI:10.1016/j.postharvbio.2014.06.013.
[16] FIGUEROA C R, OPAZO M C, VERA P, et al. Effect of postharvest treatment of calcium and auxin on cell wall composition and expression of cell wall-modifying genes in the Chilean strawberry (Fragaria chiloensis) fruit[J]. Food Chemistry, 2012, 132(4): 2014-2022. DOI:10.1016/j.foodchem.2011.12.041.
[17] BLUMENKRANTZ N, ASBOE-HANSEN G. New method for quantitative determination of uronic acids[J]. Analytical Biochemistry, 1973, 54(2): 484-489. DOI:10.1016/0003-2697(73)90377-1.
[18] REN Yuanyuan, SUN Pengpeng, WANG Xuanxuan, et al. Degradation of cell wall polysaccharides and change of related enzyme activities with fruit softening in Annona squamosa during storage[J]. Postharvest Biology and Technology, 2020, 166: 111203. DOI:10.1016/j.postharvbio.2020.111203.
[19] YOSHIDA O, NAKAGAWA H, OGURA N, et al. Effect of heat treatment on the development of polygalacturonase activity in tomato fruit during ripening[J]. Plant and Cell Physiology, 1984, 25(3): 505-509. DOI:10.1093/oxfordjournals.pcp.a076739.
[20] VICENTE A R, COSTA M L, MARTíNEZ G A, et al. Effect of heat treatments on cell wall degradation and softening in strawberry fruit[J]. Postharvest Biology and Technology, 2005, 38(3): 213-222. DOI:10.1016/j.postharvbio.2005.06.005.
[21] LIN Yifen, LIN Yuzhao, LIN Yixiong, et al. A novel chitosan alleviates pulp breakdown of harvested longan fruit by suppressing disassembly of cell wall polysaccharides[J]. Carbohydrate Polymers, 2019, 217: 126-134. DOI:10.1016/j.carbpol.2019.04.053.
[22] ALI Z M, CHIN L H, LAZAN H. A comparative study on wall degrading enzymes, pectin modifications and softening during ripening of selected tropical fruits[J]. Plant Science, 2004, 167(2): 317-327. DOI:10.1016/j.plantsci.2004.03.030.
[23] ZAHARAH S S, SINGH Z. Mode of action of nitric oxide in inhibiting ethylene biosynthesis and fruit softening during ripening and cool storage of ‘Kensington Pride’ mango[J]. Postharvest Biology and Technology, 2011, 62(3): 258-266. DOI:10.1016/j.postharvbio.2011.06.007.
[24] ZAHARAH S S, SINGH Z, SYMONS G M, et al. Mode of action of abscisic acid in triggering ethylene biosynthesis and softening during ripening in mango fruit[J]. Postharvest Biology and Technology, 2013, 75: 37-44. DOI:10.1016/j.postharvbio.2012.07.009.
[25] LIVAK K J, SCHMITTGEN T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method[J]. Methods, 2001, 25(4): 402-408. DOI:10.1006/meth.2001.1262.
[26] ZHAO Zhilei, CAO Jiankang, JIANG Weibo, et al. Maturity-related chilling tolerance in mango fruit and the antioxidant capacity involved[J]. Journal of the Science of Food and Agriculture, 2009, 89(2): 304-309. DOI:10.1002/jsfa.3443.
[27] MALUNDO T M M, SHEWFELT R L, WARE G O, et al. Sugars and acids influence flavor properties of mango (Mangifera indica)[J]. Journal of the American Society for Horticultural Science, 2001, 126(1): 115-121. DOI:10.21273/JASHS.126.1.115.
[28] 宋聪慧, 郭水欢, 史小强, 等. 褪黑素调控果蔬采后保鲜研究进展[J]. 食品科学, 2023, 44(3): 228-236. DOI:10.7506/spkx1002-6630-20220302-037
[29] 张业歆, 刘家粮, 潘永贵, 等.氯吡苯脲浸泡处理延缓采后芒果成熟和软化的生理机制[J]. 食品科学, 2021(19): 234-241. DOI:10.7506/spkx1002-6630-20200914-177.
[30] LIU Gangshuai, ZHANG Yuxin, YUN Ze, et al. Melatonin enhances cold tolerance by regulating energy and proline metabolism in litchi fruit[J]. Foods, 2020, 9(4): 454. DOI:10.3390/foods9040454.
[31] 刘丽, 林苗苗, 方金豹, 等. 秋水仙素对‘琼露’猕猴桃多倍体诱导的影响[J]. 经济林研究, 2017, 35(3): 147-151. DOI:10.14067/j.cnki.1003-8981.2017.03.021.
[32] GAWKOWSKA D, CYBULSKA J, ZDUNEK A. Structure-related gelling of pectins and linking with other natural compounds: A review[J]. Polymers, 2018, 10(7): 762. DOI:10.3390/polym10070762.
[33] 芦玉佳, 张昱, 宋美玉, 等. 外源果糖处理对采后杏果实软化的影响[J]. 食品科学, 2023, 44(11): 152-159. DOI:10.7506/spkx1002-6630-20220623-260.
[34] 郝慧慧, 邱雪, 张海红, 等. 灵武长枣贮藏过程中细胞壁降解及多糖结构的变化[J]. 中国食品学报, 2022, 22(09): 199-207. DOI:10.16429/j.1009-7848.2022.09.021.
[35] LIN Yifen, LIN Yixiong, LIN Hetong, et al. Effects of paper containing 1-MCP postharvest treatment on the disassembly of cell wall polysaccharides and softening in Younai plum fruit during storage[J]. Food Chemistry, 2018, 264: 1-8. DOI:10.1016/j.foodchem.2018.05.031.
[36] 张珅, 张翼翔, 叶洪, 等. 热处理对采后琯溪蜜柚果实汁胞粒化的影响及其与细胞壁代谢的关系[J]. 食品科学, 2021, 42(11): 17-25. DOI:10.7506/spkx1002-6630-20200714-178.
[37] JI Yaru, HU Wenzhong, LIAO Jia, et al. Ethanol vapor delays softening of postharvest blueberry by retarding cell wall degradation during cold storage and shelf life[J]. Postharvest Biology and Technology, 2021, 177: 111538. DOI:10.1016/j.postharvbio.2021.111538.
[38] 茅林春, 张上隆. 间歇低温胁迫对桃果实细胞壁代谢的影响[J]. 植物生理学报, 2001, 27(2): 151-155. DOI:10.3321/j.issn:1671-3877.2001.02.010.
[39] 罗自生, 席玙芳, 金勇丰, 等. 贮前热处理减轻柿果实冷害与细胞壁水解酶活性的关系[J]. 园艺学报, 2001, 28(6): 554-556. DOI:10.3321/j.issn:0513-353X.2001.06.013.
[40] SINGH Z, SINGH R K, SANE V A, et al. Mango-postharvest biology and biotechnology[J]. Critical Reviews in Plant Sciences, 2013, 32(4): 217-236. DOI:10.1080/07352689.2012.743399.
[41] PATTYN J, VAUGHAN-HIRSCH J, VAN DE POEL B. The regulation of ethylene biosynthesis: a complex multilevel control circuitry[J]. New Phytologist, 2021, 229(2): 770-782. DOI:10.1111/nph.16873.
[42] 杨杨, 范蓓, 申琳, 等. 芒果采后冷害发生及控制技术研究进展[J]. 食品科学, 2014, 35(07): 292-297. DOI:10.7506/spkx1002-6630-201407057.
[43] WEI Chuangqi, MA Lihua, CHENG Yudou, et al. Exogenous ethylene alleviates chilling injury of ‘Huangguan’ pear by enhancing the proline content and antioxidant activity[J]. Scientia Horticulturae, 2019, 257: 108671. DOI:10.1016/j.scienta.2019.108671.
[44] ZHU Yongchao, WANG Ke, WU Chunxia, et al. Effect of ethylene on cell wall and lipid metabolism during alleviation of postharvest chilling injury in peach[J]. Cells, 2019, 8(12): 1612. DOI:10.3390/cells8121612.
[45] SHU Pan, LI Yujing, XIANG Lanting, et al. Ethylene enhances tolerance to chilling stress in tomato fruit partially through the synergistic regulation between antioxidant enzymes and ATP synthases[J]. Postharvest Biology and Technology, 2022, 193: 112065. DOI:10.1016/j.postharvbio.2022.112065.
[46] ZHOU Ziyin, YANG Yingying, SHAN Wei, et al. Ethylene attenuates chilling injury of banana fruit via the MabHLH060/183 module in controlling phosphatidic acid formation genes[J]. Postharvest Biology and Technology, 2022, 183: 111724. DOI:10.1016/j.postharvbio.2021.111724.
[47] 赵赫, 陈受宜, 张劲松. 乙烯信号转导与植物非生物胁迫反应调控研究进展[J]. 生物技术通报, 2016, 32(10): 1-10. DOI:10.13560/j.cnki.biotech.bull.1985.2016.10.001.
[48] 史庆玲, 李忠峰, 董永彬, 等. 植物乙烯信号转导通路及其相关基因的研究进展[J]. 生物技术进展, 2019, 9(5): 449. DOI:10.19586/j.2095-2341.2019.0080.
[49] WANG Ke, YIN Xueren, ZHANG Bo, et al. Transcriptomic and metabolic analyses provide new insights into chilling injury in peach fruit[J]. Plant, Cell & Environment, 2017, 40(8): 1531-1551. DOI:10.1111/pce.12951.
[50] SHAN Shuangshuang, WANG Zhiqiang, PU Huili, et al. DNA methylation mediated by melatonin was involved in ethylene signal transmission and ripening of tomato fruit[J]. Scientia Horticulturae, 2022, 291: 110566. DOI:10.1016/j.scienta.2021.110566.
[51] KIM Y S, LEE M, LEE J H, et al. The unified ICE-CBF pathway provides a transcriptional feedback control of freezing tolerance during cold acclimation in Arabidopsis[J]. Plant Molecular Biology, 2015, 89: 187-201. DOI:10.1007/s11103-015-0365-3.
[52] HU Shunqing, MA Yuqing, XIE Bing, et al. 24-Epibrassinolide improves chilling tolerance by regulating PpCBF5-mediated membrane lipid metabolism in peach fruit[J]. Postharvest Biology and Technology, 2022, 186: 111844. DOI:10.1016/j.postharvbio.2022.111844.