Recent strategies for mitigating salinity stress in plants: A review of exogenous compound applications

Authors

  • Muchamad Imam Asrori Biotechnology Program, Faculty of Food Security, Universitas Negeri Surabaya, Surabaya 60286, Indonesia
  • Endar Hidayat Data-driven Polymer Design Group, Research Center for Macromolecules and Biomaterials, National Institute for Materials Science, Sengen, Tsukuba 305-0047, Japan

Keywords:

Salinity stress, exogenous compounds, phytohormones, antioxidants, nanoparticles

Abstract

Soil salinity is a major abiotic stress that severely limits plant growth, development, and agricultural productivity worldwide. This review summarizes recent progress in alleviating salinity stress through the external application of various compounds, including phytohormones, organic and inorganic small molecules, and nanoparticles. The application of exogenous compounds to various crop species and experimental systems consistently improves plant salt tolerance by activating convergent physiological and molecular mechanisms: (1) activation of enzyme and non-enzymatic antioxidant defense systems, reducing the ROS and MDA content; (2) maintaining ion homeostasis by limiting Na⁺ accumulation and enhancing K⁺ and Ca²⁺ uptake; (3) inducing the osmolytes; and (4) modulating the stress-responsive gene expression and hormonal crosstalk. Effectiveness depends on compound concentration, application method (seed priming, foliar spray, root absorption), plant species, and salinity severity. Synergistic combinations are often applied to enhance plant tolerance under salt stress. Future research priorities include the omics approach of regulatory networks, gene editing to validate target genes, and long-term environmental safety assessments. Practical applications provide accessible, low-cost strategies to sustain crop productivity in salt-affected regions.

References

1. Shrivastava P, Kumar R. Soil salinity: A serious environmental issue and plant growth promoting bacteria as one of the tools for its alleviation. Saudi J Biol Sci. 2015;22(2):123-31.

2. Pandit K, Chandni, Kaur S, Kumar M, Bhardwaj R, Kaur S. Chapter Six - Salinity stress: Impact on plant growth. In: Sharma A, Kumar M, Sharma P, editors. Advances in Food Security and Sustainability. 9: Elsevier; 2024. p. 145-60.

3. Hu S, Jiang P, Guo Q. Characterization of the Salt Overly Sensitive 1 (SOS1) Pathway Genes in Tea Plant (Cameliia sinensis) Under Environmental Stress. Horticulturae. 2025;11(7):855.

4. Song X, Chen J, Xu C, Cai X, Song W, Chang A, et al. Physiological and molecular mechanisms of exogenous salicylic acid in enhancing salt tolerance in tobacco seedlings by regulating antioxidant defence system and gene expression. Frontiers in Plant Science. 2025;Volume 16 - 2025.

5. Hao S, Wang Y, Yan Y, Liu Y, Wang J, Chen S. A Review on Plant Responses to Salt Stress and Their Mechanisms of Salt Resistance. Horticulturae. 2021;7(6):132.

6. Liu C, Jiang X, Yuan Z. Plant Responses and Adaptations to Salt Stress: A Review. Horticulturae. 2024;10(11):1221.

7. Athar HU, Zulfiqar F, Moosa A, Ashraf M, Zafar ZU, Zhang L, et al. Salt stress proteins in plants: An overview. Front Plant Sci. 2022;13:999058.

8. Ali A, Raddatz N, Pardo JM, Yun DJ. HKT sodium and potassium transporters in Arabidopsis thaliana and related halophyte species. Physiol Plant. 2021;171(4):546-58.

9. Sevi F, Falcinelli B, Frusciante S, Fabene E, Tosti G, D'Amato R, et al. Metabolomics of rapeseed (Brassica napus var. oleifera Del.) sprouts obtained with or without salinity from progeny seeds of mother plants grown in presence or absence of salinity. Journal of Agriculture and Food Research. 2025;19:101649.

10. Rao MJ, Duan M, Zhou C, Jiao J, Cheng P, Yang L, et al. Antioxidant Defense System in Plants: Reactive Oxygen Species Production, Signaling, and Scavenging During Abiotic Stress-Induced Oxidative Damage. Horticulturae. 2025;11(5):477.

11. Zhang H, Yu C, Zhang Q, Qiu Z, Zhang X, Hou Y, et al. Salinity survival: molecular mechanisms and adaptive strategies in plants. Front Plant Sci. 2025;16:1527952.

12. Attia H, Alamer K, Algethami B, Zorrig W, Hessini K, Gupta K, et al. Gibberellic acid interacts with salt stress on germination, growth and polyamine gene expression in fennel (Foeniculum vulgare Mill.) seedlings. Physiology and Molecular Biology of Plants. 2022;28(3):607-22.

13. Iqbal M, Ashraf M. Gibberellic acid mediated induction of salt tolerance in wheat plants: Growth, ionic partitioning, photosynthesis, yield and hormonal homeostasis. Environmental and Experimental Botany. 2013;86:76-85.

14. Álvarez-Méndez SJ, Urbano-Gálvez A, Mahouachi J. Mitigation of salt stress damages in Carica papaya L. seedlings through exogenous pretreatments of gibberellic acid and proline. Chilean journal of agricultural research. 2022;82:167-76.

15. Liu J, Meng F, Jiang A, Hou X, Liu Q, Fan H, et al. Exogenous 6-BA enhances salt tolerance of Limonium bicolor by increasing the number of salt glands. Plant Cell Reports. 2023;43(1):12.

16. Iqbal M, Ashraf M, Jamil A. Seed enhancement with cytokinins: changes in growth and grain yield in salt stressed wheat plants. Plant Growth Regulation. 2006;50(1):29-39.

17. Ma X, Zhang J, Huang B. Cytokinin-mitigation of salt-induced leaf senescence in perennial ryegrass involving the activation of antioxidant systems and ionic balance. Environmental and Experimental Botany. 2016;125:1-11.

18. da Silva TI, Gomes Dias M, Saraiva Grossi JA, Ribeiro WS, de Moraes PJ, de Araújo FF, et al. Application of phytohormones as attenuators of salt stress in Tropaeolum majus L.(Tropaeolaceae). Acta Botanica Croatica. 2022;81(1):51-60.

19. Abdel Latef AA, Tahjib-Ul-Arif M, Rhaman MS. Exogenous Auxin-Mediated Salt Stress Alleviation in Faba Bean (Vicia faba L.). Agronomy [Internet]. 2021; 11(3):[547 p.].

20. Abdel Latef AA, Akter A, Tahjib-Ul-Arif M. Foliar Application of Auxin or Cytokinin Can Confer Salinity Stress Tolerance in Vicia faba L. Agronomy [Internet]. 2021; 11(4):[790 p.].

21. Alam M, Khan MA, Imtiaz M, Khan MA, Naeem M, Shah SA, et al. Indole-3-Acetic Acid Rescues Plant Growth and Yield of Salinity Stressed Tomato (Lycopersicon esculentum L.). Gesunde Pflanzen. 2020;72(1):87-95.

22. del Pilar Cordovilla M, Aparicio C, Melendo M, Bueno M. Exogenous Application of Indol-3-Acetic Acid and Salicylic Acid Improves Tolerance to Salt Stress in Olive Plantlets (Olea europaea L. Cultivar Picual) in Growth Chamber Environments. Agronomy [Internet]. 2023; 13(3):[647 p.].

23. Ellouzi H, Ben Slimene Debez I, Amraoui S, Rabhi M, Hanana M, Alyami NM, et al. Effect of seed priming with auxin on ROS detoxification and carbohydrate metabolism and their relationship with germination and early seedling establishment in salt stressed maize. BMC Plant Biology. 2024;24(1):704.

24. Gull M, Sajid ZA, Aftab F. Alleviation of Salt Stress in Solanum tuberosum L. by Exogenous Application of Indoleacetic acid and l-Tryptophan. Journal of Plant Growth Regulation. 2023;42(5):3257-73.

25. Shiraz M, Sami F, Siddiqui H, Yusuf M, Hayat S. Interaction of Auxin and Nitric Oxide Improved Photosynthetic Efficiency and Antioxidant System of Brassica juncea Plants Under Salt Stress. Journal of Plant Growth Regulation. 2021;40(6):2379-89.

26. Srivastava A, Mahra S, Hsan N, Singh S, Sharma S, Koh J, et al. Boron nanoparticles combined with auxin alleviate salinity-induced oxidative stress in Oryza sativa L. Plant Science. 2025;359:112538.

27. Mulaudzi T, Sias G, Nkuna M, Ndou N, Hendricks K, Ikebudu V, et al. Seed Priming with MeJa Prevents Salt-Induced Growth Inhibition and Oxidative Damage in Sorghum bicolor by Inducing the Expression of Jasmonic Acid Biosynthesis Genes. International Journal of Molecular Sciences [Internet]. 2023; 24(12):[10368 p.].

28. Sheteiwy MS, Shao H, Qi W, Daly P, Sharma A, Shaghaleh H, et al. Seed priming and foliar application with jasmonic acid enhance salinity stress tolerance of soybean (Glycine max L.) seedlings. Journal of the Science of Food and Agriculture. 2021;101(5):2027-41.

29. Kasprowiak F, Wilmowicz E, Kućko A. Jasmonate-Mediated Mitigation of Salinity Stress During Germination and Early Vegetative Development in Hemp. Plants (Basel). 2025;14(18).

30. Nie R, Wu C, Ji X, Li A, Zheng X, Tang J, et al. Methyl Jasmonate Orchestrates Multi-Pathway Antioxidant Defense to Enhance Salt Stress Tolerance in Walnut (Juglans regia L.). Antioxidants. 2025; 14(8):[974 p.].

31. Labiad MH, Giménez A, Varol H, Tüzel Y, Egea-Gilabert C, Fernández JA, et al. Effect of Exogenously Applied Methyl Jasmonate on Yield and Quality of Salt-Stressed Hydroponically Grown Sea Fennel (Crithmum maritimum L.). Agronomy [Internet]. 2021; 11(6):[1083 p.].

32. Song R-F, Li T-T, Liu W-C. Jasmonic Acid Impairs Arabidopsis Seedling Salt Stress Tolerance Through MYC2-Mediated Repression of CAT2 Expression. Frontiers in Plant Science. 2021;Volume 12 - 2021.

33. Lamnai K, Anaya F, Fghire R, Zine H, Wahbi S, Loutfi K. Impact of Exogenous Application of Salicylic Acid on Growth, Water Status And Antioxidant Enzyme Activity of Strawberry Plants (Fragaria vesca L.) Under Salt Stress Conditions. Gesunde Pflanzen. 2021;73(4):465-78.

34. Ilyas M, Maqsood MF, Shahbaz M, Zulfiqar U, Ahmad K, Naz N, et al. Alleviating salinity stress in canola (Brassica napus L.) through exogenous application of salicylic acid. BMC Plant Biol. 2024;24(1):611.

35. Veloso LLdSA, Lima GSd, Silva AARd, Souza LdP, Lacerda CNd, Silva IJd, et al. Attenuation of salt stress on the physiology and production of bell peppers by treatment with salicylic acid. Semina: Ciências Agrárias. 2021;42(5):2751-68.

36. Wang Z, Dong S, Teng K, Chang Z, Zhang X. Exogenous Salicylic Acid Optimizes Photosynthesis, Antioxidant Metabolism, and Gene Expression in Perennial Ryegrass Subjected to Salt Stress. Agronomy [Internet]. 2022; 12(8):[1920 p.].

37. Ferdosi MFH, Shoaib A, Habib S, Khan KA. Modulation of salt-induced stress impact in Gladiolus grandiflorus L. by exogenous application of salicylic acid. Scientific Reports. 2021;11(1):15597.

38. Torun H, Novák O, Mikulík J, Strnad M, Ayaz FA. The Effects of Exogenous Salicylic Acid on Endogenous Phytohormone Status in Hordeum vulgare L. under Salt Stress. Plants [Internet]. 2022; 11(5):[618 p.].

39. Zhang T, Yang H. Physiological and Biochemical Mechanisms of Exogenous Calcium Chloride on Alleviating Salt Stress in Two Tartary Buckwheat (Fagopyrum tataricum) Varieties Differing in Salinity Tolerance. Phyton-International Journal of Experimental Botany. 2022;91(8).

40. Abdul Jaleel C, Manivannan P, Sankar B, Kishorekumar A, Panneerselvam R. Calcium chloride effects on salinity-induced oxidative stress, proline metabolism and indole alkaloid accumulation in Catharanthus roseus. Comptes Rendus Biologies. 2007;330(9):674-83.

41. Ren H, Zhang Y, Zhong M, Hussian J, Tang Y, Liu S, et al. Calcium signaling-mediated transcriptional reprogramming during abiotic stress response in plants. Theoretical and Applied Genetics. 2023;136(10):210.

42. Fidalgo-Illesca C, Francini A, Raffaelli A, Sebastiani L. Silicon priming triggers differential physiological, ionomic and metabolic responses in olive (Olea europaea L.) cultivars with different tolerance to salinity. Plant Physiology and Biochemistry. 2025;223:109809.

43. Peña Calzada K, Calero Hurtado A, Olivera Viciedo D, Habermann E, de Mello Prado R, de Oliveira R, et al. Regulatory Role of Silicon on Growth, Potassium Uptake, Ionic Homeostasis, Proline Accumulation, and Antioxidant Capacity of Soybean Plants Under Salt Stress. Journal of Plant Growth Regulation. 2023;42(7):4528-40.

44. Oueslati S, Ellouzi H, Jday M, Kalai FZ, Alyami NM, Hanana M, et al. Enhancing salt tolerance and growth in Brassica rapa var. rapa L. through vitamin seed priming: The role of ascorbic acid and riboflavin. Scientia Horticulturae. 2025;350:114354.

45. El-Badri AM, Batool M, Mohamed IAA, Hashem AM, Wang Z, Wang C, et al. Effective role of putrescene in mitigating harmful effects of salinity on rapeseed seedlings via modulating morpho-physiochemical and molecular processes. Industrial Crops and Products. 2025;238:122331.

46. Zulfiqar F, Moosa A, Ferrante A, Nafees M, Darras A, Nazir MM, et al. Exogenous foliar application of melatonin mitigates salt induced oxidative stress and promotes growth in Gerbera jamosonii. South African Journal of Botany. 2023;161:678-84.

47. Askari M, Hamid N, Abideen Z, Zulfiqar F, Moosa A, Nafees M, et al. Exogenous melatonin application stimulates growth, photosynthetic pigments and antioxidant potential of white beans under salinity stress. South African Journal of Botany. 2023;160:219-28.

48. Nakhaie A, Habibi G, Vaziri A. Exogenous proline enhances salt tolerance in acclimated Aloe vera by modulating photosystem II efficiency and antioxidant defense. South African Journal of Botany. 2022;147:1171-80.

49. Ramzan M, Shah AA, Ahmed MZ, Bukhari MA, Ali L, Casini R, et al. Exogenous application of glutathione and gamma amino-butyric acid alleviates salt stress through improvement in antioxidative defense system and modulation of CaXTHs stress-related genes. South African Journal of Botany. 2023;157:266-73.

50. Alharbi K, Desoky E-SM, Almuziny M, Abuzaid AO, Abu-Elsaoud AM, Algopishi UB, et al. Foliar-applied selenium nanoparticles improve antioxidant defense and photosynthetic efficiency to enhance salt stress tolerance in cowpea (Vigna unguiculata L.). Scientia Horticulturae. 2025;353:114454.

51. Elsheery NI, Nosier AM, Maswada HF, Teiba II, Elhamahmy M, Abdelrazik EM, et al. Alleviating the harmful effect of salinity on faba bean plants using selenium nanoparticles. Plant Nano Biology. 2025;12:100158.

52. Alam P, Faizan M, Sultan H, Balawi TA. Silicon oxide nanoparticles boost rice resilience to salinity by enhancing antioxidant defenses and stress regulation. Plant Science. 2025;359:112588.

53. Alam P, Yalcin M, Faizan M, Albalawi T. Response of tomato to silicon dioxide nanoparticles under salinity: Impact on photosynthesis, antioxidant enzymes activity, stress biomarkers and osmoregulatory substances. Plant Nano Biology. 2025;13:100171.

54. Mahmoud LM, Shalan AM, El-Boray MS, Vincent CI, El-Kady ME, Grosser JW, et al. Application of silicon nanoparticles enhances oxidative stress tolerance in salt stressed ‘Valencia’ sweet orange plants. Scientia Horticulturae. 2022;295:110856.

55. Li Y, Hu H, Wang R, Yin J, Zhu Y, Chen L. Silica nanoparticles promote the growth of pepper under salt stress by improving photosynthesis and modulating the water relationship. Scientia Horticulturae. 2025;350:114295.

56. Anwar T, Qureshi H, Zahid S, El-Beltagi HS, Alqahtani MD, Ismoilov I, et al. Enhancing Petroselinum crispum Tolerance to Salt Stress Through Combined Application of Green-Synthesized Silicon Nanoparticles and Melatonin. Silicon. 2025;17(18):4303-17.

57. Akhondi M, Amiri MS, Beyk-Khormizi A, Mousavi-Kouhi SM, Gautam RK, Mottaghipisheh J, et al. Zinc oxide nanoparticles as a way to improve physiology, photosynthesis, and anti-oxidative systems in Salvia leriifolia. Plant Nano Biology. 2025;14:100211.

58. Baksi S, Singh KM, Rani S, Sharma P. Melatonin-functionalized zinc oxide nanoparticles enhance salt stress tolerance in Vigna mungo L. by regulating antioxidants and ion homeostasis. Environmental Science: Nano. 2025;12(11):5056-73.

59. Mishra M, Afzal S, Yadav R, Singh NK, Zarbakhsh S. Salinity stress amelioration through selenium and zinc oxide nanoparticles in rice. Scientific Reports. 2025;15(1):27554.

60. Hamidian M, Kazemeini SA, Movahhedi Dehnavi M, Ramezanian A, Mottaghi Jahromie MR, Farsijani P, et al. Individual and combined exogenous application of melatonin and methyl Jasmonate confer salinity stress tolerance in tomato by enhancing antioxidants defense system. Scientia Horticulturae. 2025;342:114040.

61. Dogan Y, Alam P, Sultan H, Sharma R, Soysal S, Baran MF, et al. Zinc oxide nanoparticles for sustainable agriculture: A tool to combat salinity stress in rice (Oryza sativa) by modulating the nutritional profile and redox homeostasis mechanisms. Journal of Agriculture and Food Research. 2025;19:101598.

Downloads

Published

2026-04-27

How to Cite

Asrori, M. I., & Hidayat, E. (2026). Recent strategies for mitigating salinity stress in plants: A review of exogenous compound applications . Journal of Integrated Biotechnology Research , 1(1), 13–31. Retrieved from https://journal.unesa.ac.id/index.php/jibr/article/view/53013

Issue

Section

Articles
Abstract views: 72 , PDF Downloads: 59