{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T15:02:14Z","timestamp":1790002934124,"version":"4.0.1"},"reference-count":51,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T00:00:00Z","timestamp":1789603200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Salinity is one of the most harmful abiotic factors limiting crop productivity. Chitosan is a natural polysaccharide biostimulant able to alleviate salt stress, but its efficiency depends on its physicochemical properties, which remain insufficiently understood. The present study aimed to determine whether two chemically distinct forms of chitosan, i.e., high-molecular-weight chitosan (ChT) and chitosan oligosaccharide lactate (ChL), differentially mitigate the adverse effects of NaCl-induced salinity in cucumber (Cucumis sativus L.). Plants were grown hydroponically under control conditions or in the presence of 50 mmol L\u22121 NaCl, with or without medium-applied ChT or ChL (10 mg L\u22121). Salinity reduced growth and the accumulation of photosynthetic pigments, whereas chlorophyll fluorescence parameters remained largely unaffected, indicating limited impairment of PSII photochemistry. Chitosan, particularly the ChT form, partially alleviated NaCl-induced growth inhibition; ChT increased shoot and root fresh weight and leaf area under salt stress, whereas the effects of ChL were not statistically significant. Chitosan also enhanced free proline accumulation and was associated with changes in selected tricarboxylic acid (TCA) cycle intermediates, particularly malate and \u03b1-ketoglutarate, indicating an association with altered central carbon metabolism under salt stress. In contrast, neither chitosan form substantially improved the ionic status of cucumber, as excessive Na accumulation and reductions in K concentration and the K\/Na ratio persisted despite treatment. These findings suggest that the beneficial effects of chitosan, particularly ChT, in salt-stressed cucumber were associated more closely with metabolic and osmotic responses than with restoration of ion homeostasis or direct protection of the photosynthetic apparatus, with the two chitosan forms showing different response patterns under the experimental conditions used.<\/jats:p>","DOI":"10.3390\/molecules31183306","type":"journal-article","created":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T14:12:09Z","timestamp":1789999929000},"page":"3306","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Comparative Effects of High-Molecular-Weight Chitosan and Chitosan Oligosaccharide Lactate on Metabolic Adjustment and Salinity Tolerance in Cucumis sativus L."],"prefix":"10.3390","volume":"31","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2935-1491","authenticated-orcid":false,"given":"Barbara","family":"Hawrylak-Nowak","sequence":"first","affiliation":[{"name":"Department of Botany and Plant Physiology, University of Life Sciences in Lublin, Akademicka 15, 20-950 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3251-0182","authenticated-orcid":false,"given":"Maria","family":"Stasi\u0144ska-Jakubas","sequence":"additional","affiliation":[{"name":"Department of Botany and Plant Physiology, University of Life Sciences in Lublin, Akademicka 15, 20-950 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5945-1047","authenticated-orcid":false,"given":"Jan","family":"Sawicki","sequence":"additional","affiliation":[{"name":"Department of Analytical Chemistry, Medical University of Lublin, Chod\u017aki 4a, 20-093 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Renata","family":"Matraszek-Gawron","sequence":"additional","affiliation":[{"name":"Department of Botany and Plant Physiology, University of Life Sciences in Lublin, Akademicka 15, 20-950 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Weronika","family":"Jarmulska","sequence":"additional","affiliation":[{"name":"Department of Botany and Plant Physiology, University of Life Sciences in Lublin, Akademicka 15, 20-950 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Magdalena","family":"Czelej-Wrzesie\u0144","sequence":"additional","affiliation":[{"name":"Department of Vascular Surgery and Angiology, Medical University of Lublin, Staszica 16, 20-081 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4413-3748","authenticated-orcid":false,"given":"S\u0142awomir","family":"Dresler","sequence":"additional","affiliation":[{"name":"Department of Analytical Chemistry, Medical University of Lublin, Chod\u017aki 4a, 20-093 Lublin, Poland"},{"name":"Department of Plant Physiology and Biophysics, Maria Curie-Sk\u0142odowska University, Akademicka 19, 20-033 Lublin, Poland"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of salinity tolerance","volume":"59","author":"Munns","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1071\/FP09249","article-title":"Soil processes affecting crop production in salt-affected soils","volume":"37","author":"Rengasamy","year":"2010","journal-title":"Funct. Plant Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"425","DOI":"10.3109\/07388551.2014.889080","article-title":"Global plant-responding mechanisms to salt stress: Physiological and molecular levels and implications in biotechnology","volume":"35","author":"Tang","year":"2015","journal-title":"Crit. Rev. Biotechnol."},{"key":"ref_4","first-page":"100017","article-title":"Mechanisms of plant responses and adaptation to soil salinity","volume":"1","author":"Zhao","year":"2020","journal-title":"Innovation"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1007\/s12374-017-0133-9","article-title":"Salt stress tolerance; what do we learn from halophytes?","volume":"60","author":"Ali","year":"2017","journal-title":"J. Plant Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1016\/j.plaphy.2020.08.042","article-title":"Salinity induced physiological and biochemical changes in plants: An omic approach towards salt stress tolerance","volume":"156","author":"Arif","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Chen, X., Yang, J., She, D., Chen, W., Wu, J., Wang, Y., Chen, M., Li, Y., Qureshi, A.S., and Singh, A. (2025). Monitoring, reclamation and management of salt-affected lands. Water, 17.","DOI":"10.3390\/books978-3-7258-4036-6"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Bandehagh, A., and Taylor, N.L. (2020). Can alternative metabolic pathways and shunts overcome salinity induced inhibition of central carbon metabolism in crops?. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.01072"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"4804","DOI":"10.1093\/jxb\/eraf116","article-title":"Adjustments of plant primary metabolism in the face of climate change","volume":"76","author":"Bulut","year":"2025","journal-title":"J. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"119189","DOI":"10.1016\/j.carbpol.2022.119189","article-title":"Chitosan, chitosan nanoparticles and modified chitosan biomaterials, a potential tool to combat salinity stress in plants","volume":"284","author":"Balusamy","year":"2022","journal-title":"Carbohydr. Polym."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Stasi\u0144ska-Jakubas, M., and Hawrylak-Nowak, B. (2022). Protective, biostimulating, and eliciting effects of chitosan and its derivatives on crop plants. Molecules, 27.","DOI":"10.3390\/molecules27092801"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"968","DOI":"10.3390\/md8040968","article-title":"Chitosan in plant protection","volume":"8","author":"Adam","year":"2010","journal-title":"Mar. Drugs"},{"key":"ref_13","unstructured":"Hasanuzzaman, M. (2023). Chitosan for plant growth and stress tolerance. Climate-Resilient Agriculture, Vol. 2, Springer."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Huang, C., Tian, Y., Zhang, B., Hassan, M.J., Li, Z., and Zhu, Y. (2021). Chitosan (CTS) alleviates heat-induced leaf senescence in creeping bentgrass by regulating chlorophyll metabolism, antioxidant defense, and the heat shock pathway. Molecules, 26.","DOI":"10.3390\/molecules26175337"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.cropro.2005.03.010","article-title":"Chitosan as a potential natural compound to control pre and postharvest diseases of horticultural commodities","volume":"25","author":"Wilson","year":"2006","journal-title":"Crop Prot."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1007\/s13593-014-0252-3","article-title":"Chitosan antimicrobial and eliciting properties for pest control in agriculture: A review","volume":"35","author":"Xing","year":"2015","journal-title":"Agron. Sustain. Dev."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.plantsci.2013.03.007","article-title":"Multiple effects of chitosan on plant systems: Solid science or hype","volume":"208","author":"Hadwiger","year":"2013","journal-title":"Plant Sci."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Rom\u00e1n-Doval, R., Torres-Arellanes, S.P., Tenorio-Barajas, A.Y., Gomez-Sanchez, A., and Valencia-Lazcano, A.A. (2023). Chitosan: Properties and its application in agriculture in context of molecular weight. Polymers, 15.","DOI":"10.3390\/polym15132867"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"114","DOI":"10.3923\/ajps.2020.114.126","article-title":"Regulation of antioxidant system in wheat cultivars by using chitosan or salicylic acid to improve growth and yield under salinity stress","volume":"19","author":"Abdallah","year":"2020","journal-title":"Asian J. Plant Sci."},{"key":"ref_20","first-page":"342","article-title":"Alleviation of salt stress with chitosan foliar application and its effects on growth and development in tomato (Solanum lycopersicum L.)","volume":"9","year":"2022","journal-title":"Turk. J. Agric. Res."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Roy Chowdhury, M., Mehmet, M., Mukherjee, J., Debnath, A.J., and Ra\u017en\u00e1, K. (2025). Chitosan as an elicitor in plant tissue cultures: Methodological challenges. Molecules, 30.","DOI":"10.3390\/molecules30173476"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Li, L., Du, L., Cao, Q., Yang, Z., Liu, Y., Yang, H., Duan, X., and Meng, Z. (2023). Salt tolerance evaluation of cucumber germplasm under sodium chloride stress. Plants, 12.","DOI":"10.3390\/plants12162927"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Chen, C., Yu, W., Xu, X., Wang, Y., Wang, B., Xu, S., Lan, Q., and Wang, Y. (2024). Research advancements in salt tolerance of Cucurbitaceae: From salt response to molecular mechanisms. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25169051"},{"key":"ref_24","first-page":"63","article-title":"Response of cucumber plants to foliar application of chitosan and yeast under greenhouse conditions","volume":"6","author":"Shehata","year":"2012","journal-title":"Aust. J. Basic Appl. Sci."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Tan, C., Li, N., Wang, Y., Yu, X., Yang, L., Cao, R., and Ye, X. (2023). Integrated physiological and transcriptomic analyses revealed improved cold tolerance in cucumber (Cucumis sativus L.) by exogenous chitosan oligosaccharide. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24076202"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Jiao, Q., Shen, F., Fan, L., Song, Z., Zhang, J., Song, J., Fahad, S., Liu, F., Zhao, Y., and Tian, Z. (2024). Chitosan regulates the root architecture system, photosynthetic characteristics and antioxidant system contributing to salt tolerance in maize seedling. Agriculture, 14.","DOI":"10.3390\/agriculture14020304"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/s12298-018-0633-1","article-title":"Application of chitosan on plant responses with special reference to abiotic stress","volume":"25","author":"Hidangmayum","year":"2019","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1186\/s40538-026-00953-9","article-title":"Properties of chitosan and its applications as a functional material throughout the plant growth cycle and in sustainable agricultural development","volume":"13","author":"Huang","year":"2026","journal-title":"Chem. Biol. Technol. Agric."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"747","DOI":"10.35812\/CelluloseChemTechnol.2025.59.64","article-title":"Chitosan: A critical review of structural characteristics\u2013properties relationship","volume":"59","author":"Marin","year":"2025","journal-title":"Cellul. Chem. Technol."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Bigham Soostani, S., Ranjbar, M., Memarian, A., Mohammadi, M., and Yaghini, Z. (2025). Regulation of APX, SOD, and PAL genes by chitosan under salt stress in rapeseed (Brassica napus L.). BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-06815-0"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Alenazi, M.M., El-Ebidy, A.M., El-Shehaby, O.A., Seleiman, M.F., Aldhuwaib, K.J., and Abdel-Aziz, H.M.M. (2024). Chitosan and chitosan nanoparticles differentially alleviate salinity stress in Phaseolus vulgaris L. Plants, 13.","DOI":"10.3390\/plants13030398"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Elkarmout, A.F., Yang, M., and Hassan, F.A.S. (2022). Chitosan treatment effectively alleviates the adverse effects of salinity in Moringa oleifera Lam. via enhancing antioxidant system and nutrient homeostasis. Agronomy, 12.","DOI":"10.3390\/agronomy12102513"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Cao, Y., Yan, R., Sun, M., Guo, J., and Zhang, S. (2025). Effects of exogenous chitosan concentrations on photosynthesis and functional physiological traits of hibiscus under salt stress. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-06424-x"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"7024","DOI":"10.1007\/s42729-024-02021-3","article-title":"Effect of exogenous chitosan on physiological characteristics, photosynthetic parameters, and antioxidant systems of maize seedlings under salt stress","volume":"24","author":"Shen","year":"2024","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"28219","DOI":"10.1038\/s41598-025-13996-z","article-title":"Chitosan pre-and post-treatment modulates molecular and physiological responses to salinity in Brassica napus L","volume":"15","author":"Ranjbar","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Zhang, G., Wang, Y., Wu, K., Zhang, Q., Feng, Y., Miao, Y., and Yan, Z. (2021). Exogenous application of chitosan alleviates salinity stress in lettuce (Lactuca sativa L.). Horticulturae, 7.","DOI":"10.3390\/horticulturae7100342"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Geng, W., Li, Z., Hassan, M.J., and Peng, J. (2020). Chitosan regulates metabolic balance, polyamine accumulation, and Na+ transport contributing to salt tolerance in creeping bentgrass. BMC Plant Biol., 20.","DOI":"10.1186\/s12870-020-02720-w"},{"key":"ref_38","first-page":"377","article-title":"Plant secondary metabolites: Biosynthesis, classification, function and pharmacological properties","volume":"2","author":"Kabera","year":"2014","journal-title":"J. Pharm. Pharmacol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4038","DOI":"10.1093\/jxb\/erab019","article-title":"Organic acids: Versatile stress-response roles in plants","volume":"72","author":"Panchal","year":"2021","journal-title":"J. Exp. Bot."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Igamberdiev, A.U., and Eprintsev, A.T. (2016). Organic acids: The pools of fixed carbon involved in redox regulation and energy balance in higher plants. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01042"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"355","DOI":"10.30848\/PJB2020-2(24)","article-title":"Role of sugars, amino acids and organic acids in improving plant abiotic stress tolerance","volume":"52","author":"Khan","year":"2020","journal-title":"Pak. J. Bot."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1007\/s10725-010-9547-9","article-title":"Effects of citric acid as an important component of the responses to saline and alkaline stress in the halophyte Leymus chinensis (Trin.)","volume":"64","author":"Sun","year":"2011","journal-title":"Plant Growth Regul."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Tahjib-Ul-Arif, M., Zahan, M.I., Karim, M.M., Imran, S., Hunter, C.T., Islam, M.S., Mia, M.A., Hannan, M.A., Rhaman, M.S., and Hossain, M.A. (2021). Citric acid-mediated abiotic stress tolerance in plants. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22137235"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.envexpbot.2004.05.003","article-title":"Effect of various salt\u2013alkaline mixed stress conditions on sunflower seedlings and analysis of their stress factors","volume":"54","author":"Shi","year":"2005","journal-title":"Environ. Exp. Bot."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Ingrisano, R., Tosato, E., Trost, P., Gurrieri, L., and Sparla, F. (2023). Proline, cysteine and branched-chain amino acids in abiotic stress response of land plants and microalgae. Plants, 12.","DOI":"10.3390\/plants12193410"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"9348557","DOI":"10.1155\/ijog\/9348557","article-title":"Proline tagging for stress tolerance in plants","volume":"1","author":"Mushtaq","year":"2025","journal-title":"Int. J. Genom."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Chele, K.H., Steenkamp, P., Piater, L.A., Dubery, I.A., Huyser, J., and Tugizimana, F. (2021). A Global metabolic map defines the effects of a Si-based biostimulant on tomato plants under normal and saline conditions. Metabolites, 11.","DOI":"10.3390\/metabo11120820"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Quitadamo, F., De Simone, V., Beleggia, R., and Trono, D. (2021). Chitosan-induced activation of the antioxidant defense system counteracts the adverse effects of salinity in durum wheat. Plants, 10.","DOI":"10.3390\/plants10071365"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"3983","DOI":"10.1093\/jxb\/ert208","article-title":"Chlorophyll fluorescence analysis: A guide to good practice and understanding some new applications","volume":"64","author":"Murchie","year":"2013","journal-title":"J. Exp. Bot."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1042\/bst0110591","article-title":"Determination of total carotenoids and chlorophyll a and b of leaf extracts in different solvents","volume":"11","author":"Lichtenthaler","year":"1983","journal-title":"Biochem. Soc. Trans."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/BF00018060","article-title":"Rapid determination of free proline for water stress studies","volume":"39","author":"Bates","year":"1973","journal-title":"Plant Soil"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/18\/3306\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T14:15:15Z","timestamp":1790000115000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/18\/3306"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,17]]},"references-count":51,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["molecules31183306"],"URL":"https:\/\/doi.org\/10.3390\/molecules31183306","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,17]]}}}