Abstract
A two-year investigation (2023–2024) was conducted to evaluate the integrated influence of preharvest salicylic acid (SA) and maleic hydrazide (MH) treatments, alone and in combination, with a postharvest 1-methylcyclopropene (1-MCP) application on the postharvest physiology, biochemical quality and antioxidant defense mechanisms of strawberry (Fragaria × ananassa Duch.) fruits during cold storage. Results revealed that the combined preharvest application of SA + MH (T₄), further reinforced by postharvest 1-MCP treatment (T₅), markedly enhanced firmness retention, with T₅ fruits exhibiting 64–67% higher firmness than the control. This improvement was closely associated with a pronounced suppression of cell wall-degrading enzymes—polygalacturonase, pectin methylesterase, and cellulase—by 51–56% under T₅. Biochemical profiling demonstrated significant elevation in total soluble solids (TSS), titratable acidity (TA), and ascorbic acid content, accompanied by a 34–36% reduction in respiration rate under T₅, indicating delayed senescence and sustained metabolic balance.Furthermore, the combined treatments substantially enriched the phytochemical composition, with total phenolics, flavonoids, and anthocyanins increasing by 50–52%, 45–47%, and 42–43%, respectively, under T₅ compared with the control. These enhancements were paralleled by improved oxidative stress tolerance, reflected in a 47–48% reduction in malondialdehyde (MDA) content and a 40–45% upregulation in key antioxidant enzymes (SOD, CAT, APX, and POD). Principal component analysis (PCA) distinctly separated quality-related parameters (firmness, TSS, antioxidants, and phenolics) from senescence-linked traits (PG, PME, MDA, and respiration), while the correlation matrix confirmed strong positive associations among firmness, sugars, phenolics, and antioxidant enzymes. Collectively, the findings demonstrate that the integration of preharvest SA + MH with postharvest 1-MCP constitutes an effective and reproducible strategy for mitigating oxidative damage, delaying softening, and enhancing the biochemical and phytochemical quality of strawberry fruits during storage. This integrated pre- and postharvest approach offers a sustainable means to extend shelf life, preserve nutritional integrity, and improve marketable fruit quality under commercial cold storage conditions.
Data availability
Data will be available on corresponding author request.
References
Zheng, X., Zhang, J. & Hu, W. Advances in Understanding postharvest softening and quality deterioration of strawberry fruit. Trends Food Sci. Technol. 138, 422–435 (2023).
Ali, S. et al. Effect of Salicylic acid on biochemical and antioxidant attributes of Peach fruits during cold storage. Sci. Hortic. 257, 108710 (2019).
Hasanuzzaman, M., Raihan, M. R. H., Nahar, K. & Fujita, M. Role of Salicylic acid in plant abiotic stress tolerance: A critical review. Front. Plant. Sci. 13, 829982 (2022).
Mandal, S., Mallick, N. & Mitra, A. Salicylic acid-induced resistance to Fusarium oxysporum in tomato. Plant. Physiol. Biochem. 48, 443–450 (2010).
Kazemi, M., Aran, M. & Zamani, S. Effect of Salicylic acid treatments on quality characteristics of strawberry fruit during storage. Sci. Hortic. 129, 31–36 (2011).
Wang, L., He, X., Cao, Y., Wang, S. & Han, J. Exogenous Salicylic acid delays fruit softening and maintains quality of strawberry during cold storage. Postharvest Biol. Technol. 188, 111880 (2022).
García-Pastor, M. E., Serrano, M., Guillén, F., Zapata, P. J. & Valero, D. Preharvest application of maleic Hydrazide delays ripening and maintains quality in sweet Cherry. Postharvest Biol. Technol. 175, 111482 (2021).
Mahmoud, A. R., Abd El-Aziz, F. H. & Fawzi, M. I. F. Postharvest application of maleic Hydrazide improves quality and storability of Mango fruit. Sci. Hortic. 261, 108939 (2020).
Singh, V., Singh, R. & Sharma, S. Influence of maleic Hydrazide on fruit ripening and shelf life of tomato. J. Food Sci. Technol. 56, 1127–1136 (2019).
Blankenship, S. M. & Dole, J. M. 1-Methylcyclopropene: A review. Postharvest Biol. Technol. 28, 1–25 (2003).
Trainotti, L., Pavanello, A. & Casadoro, G. Different ethylene receptors show different responses during strawberry fruit development and ripening. J. Exp. Bot. 56, 2037–2046 (2005).
Perkins-Veazie, P., Collins, J. K. & Howard, L. Strawberry fruit response to 1-methylcyclopropene. Postharvest Biol. Technol. 47, 165–169 (2008).
Jiang, Y. M., Joyce, D. C. & Terry, L. A. Effects of 1-methylcyclopropene on decay and quality of strawberry fruit. Plant. Growth Regul. 35, 213–220 (2001).
Wills, R. B. H. & Ku, V. V. V. Use of 1-methylcyclopropene to extend the postharvest life of strawberries. Postharvest Biol. Technol. 26, 279–283 (2002).
Terefe, N. S., Buckow, R. & Versteeg, C. Quality-related enzymes in fruit and vegetable products: effects of novel food processing technologies. Crit. Rev. Food Sci. Nutr. 55, 902–922 (2015).
Tian, M. S. et al. Responses of strawberry fruit to 1-methylcyclopropene. Postharvest Biol. Technol. 19, 153–160 (2000).
Singleton, V. L. & Rossi, J. A. Colorimetry of total phenolics with phosphomolybdic–phosphotungstic acid reagents. Am. J. Enol. Vitic. 16, 144–158 (1965).
Chang, C., Yang, M., Wen, H. & Chern, J. Estimation of total flavonoid content in propolis by two complementary colorimetric methods. J. Food Drug Anal. 10, 178–182 (2002).
Giampieri, F. et al. The strawberry: Composition, nutritional quality, and impact on human health. Food Funct. 5, 1271–1291 (2014).
Davey, M. W. et al. Plant L-ascorbic acid: Chemistry, function, metabolism, bioavailability and effects of processing. J. Sci. Food Agric. 80, 825–860 (2000).
Hodges, D. M., DeLong, J. M., Forney, C. F. & Prange, R. K. Improving the thiobarbituric acid-reactive substances assay for estimating lipid peroxidation in plant tissues. Physiol. Plant. 107, 336–341 (1999).
Beauchamp, C. & Fridovich, I. Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal. Biochem. 44, 276–287 (1971).
Aebi, H. Catalase in vitro. Methods Enzymol. 105, 121–126 (1984).
Nakano, Y. & Asada, K. Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant. Cell. Physiol. 22, 867–880 (1981).
Chance, B. & Maehly, A. C. Assay of catalases and peroxidases. Methods Enzymol. 2, 764–775 (1955).
FAO/WHO. Pesticide residues in food—Maleic hydrazide. FAO Plant Protection Paper. (1997).
United States Environmental Protection Agency (USEPA). Maleic hydrazide; exemption from tolerance. Fed Regist (2023).
De Bruno, N., Rinaldi, M. & Moccia, S. Postharvest quality and shelf life of strawberry fruit under cold storage. Int. J. Food Sci. Technol. 60, 123–134 (2023).
Qaderi, R. et al. Effect of storage conditions on quality and bioactive compounds of strawberry fruit. Acta Hortic. 1364, 409–416 (2023).
Pols, S., Williams, E. & de Vries, F. Strategies for extending strawberry shelf life. Acta Hortic. 1256, 639–646 (2019).
Brummell, D. A. & Harpster, M. H. Cell wall metabolism in fruit softening and quality maintenance. Plant. Mol. Biol. 47, 311–340 (2001).
Langer, S. E., Almenar, E. & Aday, M. S. 1-Methylcyclopropene reduces cell wall degrading enzyme activity in strawberries. Postharvest Biol. Technol. 186, 111860 (2022).
Win, N. M., Yoo, Y. & Kang, I. K. Synergistic effects of Salicylic acid and 1-MCP on enzyme activity and fruit quality of strawberry. Postharvest Biol. Technol. 154, 95–104 (2019).
Yoo, Y., Kim, J. H., Win, N. M. & Kang, I. K. Ethylene Inhibition delays strawberry fruit senescence. Postharvest Biol. Technol. 194, 112068 (2022).
Li, M. et al. Salicylic acid represses cell wall hydrolases and delays strawberry softening. Sci. Hortic. 316, 112013 (2023).
Aghdam, M. S., Yousefi, S., Fard, J. R. & Asghari, M. Synergistic effects of Salicylic acid and maleic Hydrazide on postharvest quality of strawberry. Postharvest Biol. Technol. 212, 113126 (2024).
Taiz, L. & Zeiger, E. Plant Physiology 3rd edn (Sinauer Associates, 2002).
Zhu, Y. et al. Molecular mechanisms of maleic hydrazide-mediated growth regulation in plants. Biotechnol. Biofuels. 14, 99 (2021).
Chen, X. et al. 1-Methylcyclopropene delays senescence and reduces respiration in strawberry fruit. Food Chem. 345, 128853 (2021).
Meng, X. et al. Salicylic acid enhances antioxidant defense and delays senescence in strawberry fruit. Antioxidants 12, 501 (2023).
Zhang, L., Wang, R., Cao, Y. & Li, S. Salicylic acid enhances phenylpropanoid metabolism and antioxidant capacity in strawberry fruit. Food Chem. 414, 135723 (2023).
Acknowledgements
The Deanship of Scientific Research (DSR) at King Abdulaziz University (KAU) has funded this project, under grant no. (RG- 6- 130-43).
Author information
Authors and Affiliations
Contributions
Neha Sharma: Data curation; Formal analysis; Investigation; Methodology; Project administration; Resources; Software; Supervision; Validation; Visualization; Roles/Writing – original draft; Parshant Bakshi : Data curation; Formal analysis; Investigation; Methodology; Project administration; Resources; Software; Supervision; Validation; Visualization; Roles/Writing – original draft; Writing – review & editing. Zahid Nabi Sheikh: Conceptualization; Data curation; Formal analysis; Methodology; Project administration, Roles/Writing – original draft; Writing – review & editing; Balbir Dhotra : Roles/Writing – original draft; Writing – review & editing. Hesham F. Alharby : Supervision; Validation; Visualization. Funding acquisition and Othman R. Alzahrani : Supervision; Validation; Visualization. Funding acquisition and Khalid Rehman Hakeem : Supervision; Validation; Visualization., Mehdi Rahimi: Formal analysis; Project administration, Writing – review & editing.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Plant source
Experimental material was collected from the Research field of the Division of Fruit Science, Sher-e-Kashmir University of Agricultural Sciences and Technology of Jammu (SKUAST-J), Chatha, India (32.63°N, 74.85°E; 332 m above sea level). The site experiences a subtropical climate with mild winters and warm summers. Uniform, healthy, and disease-free plants of Fragaria × ananassa Duch. cv. ‘Nabila’ grown under open-field conditions in raised beds were selected. Standard agronomic practices were maintained throughout the crop cycle.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
About this article
Cite this article
Sharma, N., Bakshi, P., Dhotra, B. et al. Integrated preharvest salicylic acid maleic hydrazide and postharvest 1 MCP delay softening and preserve strawberry quality during cold storage. Sci Rep (2026). https://doi.org/10.1038/s41598-026-36487-1
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-026-36487-1