Plant Soil Environ., 2026, 72(1):39-48 | DOI: 10.17221/374/2025-PSE

The changes in growth and metabolic adaptation responses in Java plum seedlings exposed to Cassia javanica extract under salinityOriginal Paper

Abeer H. Elhakem ORCID...1, Rasha S. El-Serafy ORCID...2
1 Department of Biology, College of Sciences and Humanities, Prince Sattam Bin Abdulaziz University, Al-Kharj, Saudi Arabia
2 Horticulture Department, Faculty of Agriculture, Tanta University, Tanta, Egypt

Developing and employing new, sustainable, and eco-friendly biostimulants that enhance plant growth and alleviate the harmful effects of environmental challenges is a major focus for many researchers. Salt stress is a critical constraint on plant growth and a limiting factor in crop productivity, particularly during the early developmental stages in the nurseries. Syzygium cumini (L.) Skeels (Java plum) is an important fruit tree and widely cultivated in gardens as an ornamental plant. This study was designed to develop Cassia javanica subsp. nodosa leaf extract (CLE) as a new sustainable and eco-friendly biostimulant capable of triggering the metabolic adaptation to salt stress in Java plum seedlings grown in nurseries. CLE successfully mitigated reductions in growth, biomass yield, and secondary metabolite production caused by salinity. Although salt stress depressed morphological characters and biomass yield, CLE foliar spray enhanced these parameters. Moreover, CLE enhanced the ferric reducing antioxidant potential, catalase, and superoxide dismutase enzyme activities, increased phenolic content, and reduced hydrogen peroxide (H2O2) accumulation and lipid peroxidation. Additionally, CLE application increased seedling biomass and stimulated antioxidant activity, osmoprotectant accumulation, and overall tolerance to salinity stress. These observations provide new insights into CLE’s potential as an eco-friendly biostimulant for enhancing salt tolerance in Java plum seedlings.

Keywords: abiotic stress; toxicity; osmotic stress; nursery application; osmoregulation; sustainable biostimulants

Received: August 26, 2025; Revised: January 12, 2026; Accepted: January 13, 2026; Prepublished online: January 13, 2026; Published: January 29, 2026  Show citation

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Elhakem AH, El-Serafy RS. The changes in growth and metabolic adaptation responses in Java plum seedlings exposed to Cassia javanica extract under salinity. Plant Soil Environ. 2026;72(1):39-48. doi: 10.17221/374/2025-PSE.
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    References

    1. Ábrahám E., Hourton-Cabassa C., Erdei L., Szabados L. (2010): Methods for determination of proline in plants. In: Sunkar R. (ed.): Plant Stress Tolerance: Methods and Protocols. Totowa, Humana Press, 317-331. Go to original source...
    2. Acosta-Motos J.R., Ortuño M.F., Bernal-Vicente A., Diaz-Vivancos P., Sanchez Blanco M.J., Hernandez J.A. (2017): Plant responses to salt stress: adaptive mechanisms. Agronomy, 7: 18. Go to original source...
    3. Aebi H. (1984): Catalase in vitro. Methods in Enzymology, 105: 121-126. Go to original source... Go to PubMed...
    4. Ahmad P., Nabi G., Jeleel C.A., Umar S. (2011): Free radical production, oxidative damage and antioxidant defense mechanisms in plants under abiotic stress. In: Ahmad P., Umar S. (eds.): Oxidative Stress: Role of Antioxidants in Plants. New Delhi, Studium Press, 19-53.
    5. Ahmad P., Sharma S. (2008): Salt stress and phyto-biochemical responses of plants. Plant, Soil and Environment, 54: 89-99. Go to original source...
    6. Alayafi A.H., Dahab A.A., El-Sheshtawy A.A., Sharma A., Elhakem A., Youssef S.M., El-Serafy R.S. (2025): Stimulatory effect of Delo-nix regia flower extract in protecting Syzygium cumini seedlings from salinity. Plants, 14: 875. Go to original source...
    7. Al-Menaie H.S., Al-Ragam O., Al-Shatti A., Mathew M., Suresh N. (2010): The effects of different treatments on seed germination of the Cassia fistula L. and Cassia nodosa Buch.-Ham. ex Roxb. in Kuwait. African Journal of Agricultural Research, 5: 230-235.
    8. Al-Saif A.M., Ahmed M.E.M., Taha M.A., Sharma A., El-Sheshtawy A.A., Abouelsaad I.A., El-Serafy R.S., Mahdy R.M. (2024): Pre-harvest applications improve the postharvest storage and quality of tomato fruits by enhancing the nutritional value and antioxidant system. Horticulturae, 10: 1248. Go to original source...
    9. Alshallash K.S., Mohamed M.F., Dahab A.A., Abd El-Salam H.S., El-Serafy R.S. (2022): Biostimulation of Plectranthus amboinicus (Lour.) spreng. with different yeast strains: morphological performance, productivity, phenotypic plasticity, and antioxidant activity. Horticulturae, 8: 887. Go to original source...
    10. AOAC (1995): Official Method of Analysis. 16th Edition. Arlington, Association of Official Analytical Chemists International.
    11. Arya V., Yadav J.P. (2011): Comparative assessment of relative antioxidant activity of sequential leaf extracts of Cassia occidentalis L. and Cassia tora L. Pharmacologyonline, 1: 529-543.
    12. Atteya A.K.G., Albalawi A.N., El-Serafy R.S., Albalawi K.N., Bayomy H.M., Genaidy E.A.E. (2021): Response of Moringa oleifera seeds and fixed oil production to vermicompost and NPK fertilizers under calcareous soil conditions. Plants, 10: 1998. Go to original source...
    13. Atteya A.K.G., El-Serafy R.S., El-Zabalawy K.M., Elhakem A., Genaidy E.A.E. (2022a): Brassinolide maximized the fruit and oil yield, induced the secondary metabolites, and stimulated linoleic acid synthesis of Opuntia ficus-indica oil. Horticulturae, 8: 452. Go to original source...
    14. Atteya A.K.G., El-Serafy R.S., El-Zabalawy K.M., Elhakem A., Genaidy E.A.E. (2022b): Exogenously supplemented proline and phenyl-alanine improve growth, productivity, and oil composition of salted moringa by upregulating osmoprotectants and stimulating anti-oxidant machinery. Plants, 11: 1553. Go to original source... Go to PubMed...
    15. Attia M.A.E.H., El-Enin M.M.A., Tahoun A.M.A., Abdelghany F.I.M., El-Serafy R.S. (2022): Productivity of some barley cultivars as affected by supplemental irrigation under rainfed conditions. Australian Journal of Crop Science, 16: 665-667. Go to original source...
    16. Ayyanar M., Subash-Babu P. (2012): Syzygium cumini (L.) Skeels: a review of its phytochemical constituents and traditional uses. Asian Pacific Journal of Tropical Biomedicine, 2: 240-246. Go to original source...
    17. Bahgat A.R., Dahab A.A., Elhakem A., Gururani M.A., El-Serafy R.S. (2023): Integrated action of rhizobacteria with Aloe vera and moringa leaf extracts improves defense mechanisms in Hibiscus sabdariffa L. cultivated in saline soil. Plants, 12: 3684. Go to original source... Go to PubMed...
    18. Belmain S., Neal G., Ray D., Golob P. (2001): Insecticidal and vertebrate toxicity associated with ethnobotanicals used as post-harvest protectants in Ghana. Food and Chemical Toxicology, 39: 287-291. Go to original source... Go to PubMed...
    19. Benzie I.F., Strain J.J. (1996): The ferric reducing ability of plasma (FRAP) as a measure of "antioxidant power": the FRAP assay. Annals of Biochemistry, 239: 70-76. Go to original source... Go to PubMed...
    20. Bi B., Tang J., Han S., Guo J., Miao Y. (2017): Sinapic acid or its derivatives interfere with abscisic acid homeostasis during Arabidopsis thaliana seed germination. BMC Plant Biology, 17: 1-12. Go to original source... Go to PubMed...
    21. Dere S., Günes T., Sivaci R. (1998): Spectrophotometric determination of chlorophyll-A, B and total carotenoid contents of some algae species using different solvents. Turkish Journal of Botany, 22: 13-17.
    22. Dewanto V., Wu X., Adom K.K., Liu R.H. (2002): Thermal processing enhances the nutritional value of tomatoes by increasing total antioxidant activity. Journal of Agriculture and Food Chemistry, 50: 3010-3014. Go to original source... Go to PubMed...
    23. El-Nagar A.H., Ghanem K.Z., Hassan F.A.S., Fetouh M.I., El-Serafy R.S., Moussa M.M. (2025): The changes in growth, yield, and biologically active compounds of essential oil in Trachyspermum ammi L. upon rhizobacteria and seaweed applications. Plant, Soil and Environment, 71: 565-580. Go to original source...
    24. El-Serafy R.S. (2020): Phenotypic plasticity, biomass allocation, and biochemical analysis of cordyline seedlings in response to oligochi-tosan foliar spray. Journal of Soil Science and Plant Nutrition, 20: 1503-1514. Go to original source...
    25. El-Serafy R.S., Dahab A.A., Ghanem K.Z., Elhakem A., Bahgat A.R., Venkatesh J., El-Sheshtawy A.A., Badawy A. (2024): As a natural antioxidant: Sesbania grandiflora leaf extract enhanced growth and yield performance, active ingredients and tolerance of Hibiscus sabdariffa L. under salt-affected soil. Journal of Soil Science and Plant Nutrition, 24: 3406-3420. Go to original source...
    26. El-Serafy R.S., El-Sheshtawy A.A., Atteya A.K., Al-Hashimi A., Abbasi A.M., Al-Ashkar I. (2021): Seed priming with silicon as a potential to increase salt stress tolerance in Lathyrus odoratus. Plants, 10: 2140. Go to original source... Go to PubMed...
    27. El-Serafy R.S., El-Sheshtawy A.A., Dahab A.A. (2023): Fruit peel soil supplementation induces physiological and biochemical toler-ance in Schefflera arboricola L. grown under heat conditions. Journal of Soil Science and Plant Nutrition, 23: 1046-1059. Go to original source...
    28. Eraslan F., Inal A., Savasturk O., Gunes A. (2007): Changes in antioxidative system and membrane damage of lettuce in response to salinity and boron toxicity. Scientia Horticulturae, 114: 5-10. Go to original source...
    29. Estefan G., Sommer R., Ryan J. (2013): Methods of Soil, Plant, and Water Analysis: A Manual for the West Asia and North Africa Region. 3rd Edition. Beirut, International Center for Agricultural Research in the Dry Areas, 84-105.
    30. Gémes K., Kim Y.J., Park K.Y., Moschou P.N., Andronis E., Valassaki C., Roussis A., Roubelakis-Angelakis K.A. (2016): An NADPH-oxidase/polyamine oxidase feedback loop controls oxidative burst under salinity. Plant Physiology, 172: 1418-1431. Go to original source... Go to PubMed...
    31. Ghanem K.Z., Hasham M.M., El-Sheshtawy A.A., El-Serafy R.S., Sheta M.H. (2022): Biochar stimulated actual evapotranspiration and wheat productivity under water deficit conditions in sandy soil based on non-weighing lysimeter. Plants, 11: 3346. Go to original source... Go to PubMed...
    32. Giannopolitis C.N., Ries S.K. (1977): Superoxide dismutases: I. Occurrence in higher plants. Plant Physiology, 59: 309-314. Go to original source... Go to PubMed...
    33. Gururani M.A., Atteya A.K., Elhakem A., El-Sheshtawy A.A., El-Serafy R.S. (2023): Essential oils prolonged the cut carnation longevi-ty by limiting the xylem blockage and enhancing the physiological and biochemical levels. PLoS One, 3: e0281717. Go to original source... Go to PubMed...
    34. Hadwan M.H. (2018): Simple spectrophotometric assay for measuring catalase activity in biological tissues. BMC Biochemistry, 19: 7. Go to original source... Go to PubMed...
    35. Kaur G., Sharma A., Guruprasad K., Pati P.K. (2014): Versatile roles of plant NADPH oxidases and emerging concepts. Biotechnology Advances, 32: 551-563. Go to original source... Go to PubMed...
    36. Kaur P., Arora S. (2011): Superoxide anion radical scavenging activity of Cassia siamea and Cassia javanica. Medicinal Chemistry Research, 20: 9-15. Go to original source...
    37. Khurm M., Wang X., Zhang H., Hussain S.N., Qaisar M.N., Hayat K., Saqib F., Zhang X., Zhan G., Guo Z. (2021): The genus Cassia L.: ethnopharmacological and phytochemical overview. Phytotherapy Research: PTR, 35: 2336-2385. Go to original source...
    38. König S., Feussner K., Kaever A., Landesfeind M., Thurow C., Karlovsky P., Gatz C., Polle A., Feussner I. (2014): Soluble phenylpro-panoids are involved in the defense response of Arabidopsis against Verticillium longisporum. New Phytologist, 202: 823-837. Go to original source... Go to PubMed...
    39. Kumar A., Singh S., Gaurav A.K., Srivastava S., Verma J.P. (2020): Plant growth-promoting bacteria: biological tools for the mitigation of salinity stress in plants. Frontiers in Microbiology, 11: 1216. Go to original source... Go to PubMed...
    40. Kumavat U.C., Gopalkrishnan B. (2024): Differentiation of two allied pink Cassia species based on physicochemical and chemotaxo-nomic studies. World Journal of Pharmacological Science Research, 3: 113-121.
    41. Landi M. (2017): Commentary to: "Improving the thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds" by Hodges et al., Planta (1999) 207: 604-611. Planta, 245: 1067. Go to original source...
    42. Lavanya B., Maheswaran A., Vimal N., Vignesh K., Uvarani K.Y., Varsha R. (2018): An overall view of Cassia species phytochemical constituents and its pharmacological uses. International Journal of Pharmacology Science Research, 3: 2455-4685.
    43. Leyva A., Quintana A., Sánchez M., Rodríguez E.N., Cremata J., Sánchez J.C. (2008): Rapid and sensitive anthrone-sulfuric acid assay in microplate format to quantify carbohydrate in biopharmaceutical products: method development and validation. Biologicals, 36: 134-141. Go to original source... Go to PubMed...
    44. Liang X., Zhang L., Natarajan S.K., Becker D.F. (2013): Proline mechanisms of stress survival. Antioxidants and Redox Signalling, 19: 998-1011. Go to original source...
    45. Liu S.Y., Liao C.K., Lo C.T., Yang H.H., Lin K.C., Peng K.C. (2016): Chrysophanol is involved in the biofertilization and biocontrol activities of Trichoderma. Physiological and Molecular Plant Pathology, 96: 1-7. Go to original source...
    46. Mahdy R.M., Al-Saif A.M., Ahmed M.E.M., Abd El-Bary T.S., Sharma A., El-Sheshtawy A.A., El-Serafy R.S., Abd El-Ghany T.S. (2024): Evaluation of two different methods of fulvic acid application (seed priming and foliar spray) on growth, yield, and nutri-tional quality of pea (Pisum sativum L.). Plants, 13: 3380. Go to original source... Go to PubMed...
    47. Mariotti F., Tomé D., Mirand P.P. (2008): Converting nitrogen into protein - beyond 6.25 and Jones' factors. Critical Reviews in Food Science and Nutrition, 48: 177-184. Go to original source...
    48. Pardo J.M. (2010): Biotechnology of water and salinity stress tolerance. Current Opinion Biotechnology, 21: 185-196. Go to original source... Go to PubMed...
    49. Patterson B.D., Macrae E.A., Ferguson I.B. (1984): Estimation of hydrogen peroxide in plant extracts using titanium (IV). Analytical Biochemistry, 139: 487-492. Go to original source... Go to PubMed...
    50. Paul D., Lade H. (2014): Plant-growth-promoting rhizobacteria to improve crop growth in saline soils: a review. Agronomy for Sus-tainable Development, 34: 730-752. Go to original source...
    51. Prateeksha Yusuf M.A., Singh B.N., Sudheer S., Kharwar R.N., Siddiqui S., Abdel-Azeem A.M., Fraceto L.F., Dashora K., Gupta V.K. (2019): Chrysophanol: a natural anthraquinone with multifaceted biotherapeutic potential. Biomolecules, 9: 68. Go to original source...
    52. Rasool S., Hameed A., Azooz M.M., Siddiqi T.O., Ahmad P. (2013): Salt stress: causes, types and responses of plants. In: Ahmad P., Azooz M.M., Prasad M.N.V. (eds.): Ecophysiology and Responses of Plants under Salt Stress. New York, Springer, 1-24. Go to original source...
    53. Sarvade S., Gautam D.S., Bhalawe S., Bisen P.K. (2016): An overview of potential multipurpose agroforestry tree species, Syzygium cuminii (L.) Skeels in India. Journal of Applied Natural Science, 8: 1714-1719. Go to original source...
    54. Shafi A., Zahoor I., Mushtaq U. (2019): Proline accumulation and oxidative stress: diverse roles and mechanism of tolerance and adap-tation under salinity stress. In: Akhtar M.S. (ed.): Salt Stress, Microbes, and Plant Interactions: Mechanisms and Molecular Ap-proaches 2. Singapore, Springer Nature Singapore Pte Ltd., 269-300. Go to original source...
    55. Shahidi F., Naczk M. (2004): Cereals, legumes, and nuts. In: Kakade A. (ed.): Phenolics in Food and Nutraceuticals. Boca Raton, CRC Press, 117-166. Go to original source...
    56. Sheha A.M., Abou El-Enin M.M., El-Hashash E.F., Rady M.M., El-Serafy R.S., Shaaban A. (2023): The productivity and overall bene-fits of faba bean-sugar beet intercropping systems interacted with foliar applied nutrients. Journal of Plant Nutrition, 46: 1683-1700. Go to original source...
    57. Sheta M.H., Abd El-Wahed A.H.M., Elshaer M.A., Bayomy H.M., Ozaybi N.A., Abd-Elraheem M.A.M., El-Sheshtawy A.A., El-Serafy R.S., Moustafa M.M.I. (2024): Green synthesis of zinc and iron nanoparticles using Psidium guajava leaf extract stimulates cowpea growth, yield, and tolerance to saline water irrigation. Horticulturae, 10: 915. Go to original source...
    58. Tewari S.K., Singh D., Nainwal R.C. (2017): Horticultural management of Syzygium cumini: Syzygium cumini and other underutilized species. In: Nair K.N. (ed.): The Genus Syzygium. Boca Raton, CRC Press, 215-236. Go to original source...
    59. Vogt T. (2010): Phenylpropanoid biosynthesis. Molecule Plant, 3: 2-20. Go to original source... Go to PubMed...
    60. Youssef S.M., El-Serafy R.S., Ghanem K.Z., Elhakem A., Abdel Aal A.A. (2022): Foliar spray or soil drench: microalgae application impacts on soil microbiology, morpho-physiological and biochemical responses, oil and fatty acid profiles of Chia plants under alka-line stress. Biology, 11: 1844. Go to original source... Go to PubMed...
    61. Zafar Z., Rasheed F., Atif R.M., Maqsood M., Gailing O. (2021): Salicylic acid-induced morpho-physiological and biochemical changes triggered water deficit tolerance in Syzygium cumini L. saplings. Forests, 12: 491. Go to original source...
    62. Zali A.G., Ehsanzadeh P. (2018): Exogenously applied proline as a tool to enhance water use efficiency: case of fennel. Agriculture Water Management, 197: 138-146. Go to original source...

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