Salt Tolerance Mechanisms and Strategies for Enhancing Crop Productivity in Vegetable Crops: A Review

Abbas, F, Faried, H N, Akhtar, G, Ullah, S, Javed, T, Shehzad, M A, and Abeed, A H (2023) Cucumber grafting on indigenous cucurbit landraces confers salt tolerance and improves fruit yield by enhancing morpho-physio-biochemical and ionic attributes. Scientific Reports, 13(1):21697.

Abbas, W, Ashraf, M, and Akram, N A (2010) Alleviation of salt-induced adverse effects in eggplant (Solanum melongena L.) by glycinebetaine and sugarbeet extracts. Sci. Hortic., 125:188–195.

Abd El-Fattah, B E, Haridy, A G, and Abbas, H S (2020) Response to planting date, stress tolerance and genetic diversity analysis among okra (Abelmoschus esculentus (L.) Moench.) varieties. Genetic Resources and Crop Evolution, 67:831-851.

Abdel-Moneam MA, Seadh SE, El-Sherpiny M, Mohamed AI (2024) Effect of irrigation and spraying with boron and potassium silicate in improving the quality and productivity of sugar beet under water stress conditions. Iraqi Journal of Desert Studies 14(2).

Abdullah Z, Ahmad R (1982) Salt tolerance of Solanum tuberosum L. growing on saline soils amended with gypsum.

Abobatta, W F (2020) Plant responses and tolerance to combined salt and drought stress. In Salt and Drought Stress Tolerance in Plants: Signaling Networks and Adaptive Mechanisms, pp. 17-52.

Acosta-Motos, J R, Ortuno, M F, Bernal-Vicente, A, Diaz-Vivancos, P, Sanchez-Blanco, M J, and Hernandez, J A (2017) Plant responses to salt stress: adaptive mechanisms. Agronomy, 7(1):18.

Adhikari, N D, Simko, I, and Mou, B (2019) Phenomic and physiological analysis of salinity effects on lettuce. Sensors, 19(21):4814.

Afzal, M, Hindawi, S E S, Alghamdi, S S, Migdadi, H H, Khan, M A, Hasnain, M U, and Sohaib, M (2023) Potential breeding strategies for improving salt tolerance in crop plants. Journal of Plant Growth Regulation, 42(6):3365-3387.

Ahmad, P and Jhon, R (2005) Effect of salt stress on growth and biochemical parameters of Pisum sativum L. Arch. Agron. Soil Sci., 51:665–672.

Al-Gaadi KA, Tola E, Madugundu R, Zeyada AM, Alameen AA, Edrris MK, et al. (2024) Response of leaf photosynthesis, chlorophyll content and yield of hydroponic tomatoes to different water salinity levels. PLOS One 19(2):e0293098.

Albacete, A, Martínez-Andujar, C, Ghanem, M E, Acosta, M, Sánchez-Bravo, J, Asins, M J, Cuartero, J, Lutts, S, Dodd, I C, and Pérez-Alfocea, F (2009) Rootstock-mediated changes in xylem ionic and hormonal status are correlated with delayed leaf senescence and increased leaf area and crop productivity in salinised tomato. Plant Cell Env., 32:928–938.

Alharbi, K, Al-Osaimi, A A, and Alghamdi, B A (2022) Sodium chloride (NaCl)-induced physiological alteration and oxidative stress generation in Pisum sativum (L.): A toxicity assessment. ACS Omega, 7(24):20819-20832.

Ali N, Rafiq R, Wijaya L, Ahmad A, Kaushik P (2024) Exogenous citric acid improves growth and yield by concerted modulation of antioxidant defense system in brinjal (Solanum melongena L.) under salt-stress. Journal of King Saud University-Science 36(1):103012.

Ali, M R (2020) Salinity tolerance of four winter vegetables. Master's thesis, Sher-e-Bangla Agricultural University, Dhaka, Bangladesh.

Ali, M, Kamran, M, Abbasi, G H, Saleem, M H, Ahmad, S, Parveen, A, and Fahad, S (2021) Melatonin-induced salinity tolerance by ameliorating osmotic and oxidative stress in the seedlings of two tomato (Solanum lycopersicum L.) cultivars. Journal of Plant Growth Regulation, 40:2236-2248.

Ali, M, Niaz, Y, Abbasi, G H, Ahmad, S, Malik, Z, Kamran, M, and Ayaz, M (2021) Exogenous zinc induced NaCl tolerance in okra (Abelmoschus esculentus) by ameliorating osmotic stress and oxidative metabolism. Communications in Soil Science and Plant Analysis, 52(7):743-755.

Alkhatib, R, Abdo, N, and Mheidat, M (2021) Photosynthetic and ultrastructural properties of eggplant (Solanum melongena) under salinity stress. Horticulturae, 7(7):181.

Alo, Sora S (2019) Germination, Growth and Yield Responses of Released Tomato (Solanum lycopersicum L.) Varieties to Salt Stress. Doctoral dissertation, Jimma University.

Alqardaeai, T, Alharbi, A, Alenazi, M, Alomran, A, Elfeky, A, Osman, M, and Pailles, Y (2024) Effect of Tomato Grafting onto Novel and Commercial Rootstocks on Improved Salinity Tolerance and Enhanced Growth, Physiology, and Yield in Soilless Culture.

Altuntas, O, Dasgan, H Y, Akhoundnejad, Y, and Nas, Y (2024) Unlocking the potential of pepper plants under salt stress: Mycorrhizal effects on physiological parameters related to plant growth and gas exchange across tolerant and sensitive genotypes. Plants, 13(10):1380.

Alvan, H A, Jabbarzadeh, Z, Fard, J R, and Noruzi, P (2025) Selenium foliar application alleviates salinity stress in sweet william (Dianthus barbatus L.) by enhancing growth and reducing oxidative damage. Scientific Reports, 15(1):5570.

Alzubaidy, H S (2020) Induction of salt tolerance by Enterobacter sp. SA187 in the model organism Arabidopsis thaliana.

Aman Shamil, N (2022) Role of exogenous application of proline and glycine betaine in the salinity tolerance of Solanaceae family: a review. Acta Scientific AGRICULTURE, 6(9).

Andrade, A, Masciarelli, O, Alemano, S, Luna, V, and Abdala, G (2008) Tomato mutants sensitive to abiotic stress display different abscisic acid content and metabolism during germination. Seed Science Research, 18:249-258.

Anjum S, Ain QU, Sarwar M, Alam MW, Mushtaq Z, Mukhtar A, et al. (2025) Chitosan-induced modification in morpho-physiological, biochemical, and yield attributes of pea (Pisum sativum L.) under salt stress. Journal of Soil Science and Plant Nutrition 1-11.

Arif, Y, Singh, P, Siddiqui, H, Bajguz, A, and Hayat, S (2020) Salinity induced physiological and biochemical changes in plants: An omic approach towards salt stress tolerance. Plant Physiology and Biochemistry, 156:64-77.

Ashraf, M, and McNeilly, T (2004) Salinity tolerance in Brassica oilseeds. Critical Reviews in Plant Sciences, 23(2):157-174.

Ashraf, M, Arfan, M, and Ahmad, A (2003) Salt tolerance in okra: Ion relations and gas exchange characteristics. J. Plant Nutr., 26:63–79.

Aslam, S, Gul, N, Mir, M A, Asgher, M, Al-Sulami, N, Abulfaraj, A A, and Qari, S (2021) Role of jasmonates, calcium, and glutathione in plants to combat abiotic stresses through precise signaling cascade. Frontiers in Plant Science, 12:668029.

Assaha, D V, Ueda, A, Saneoka, H, Al-Yahyai, R, and Yaish, M W (2017) The role of Na+ and K+ transporters in salt stress adaptation in glycophytes. Frontiers in Physiology, 8:509.

Atieno, J, Li, Y, Langridge, P, Dowling, K, Brien, C, Berger, B, and Sutton, T (2017) Exploring genetic variation for salinity tolerance in chickpea using image-based phenotyping. Scientific Reports, 7(1):1300.

Atta, K, Mondal, S, Gorai, S, Singh, A P, Kumari, A, Ghosh, T, and Jespersen, D (2023) Impacts of salinity stress on crop plants: Improving salt tolerance through genetic and molecular dissection. Frontiers in Plant Science, 14:1241736.

Ayub, M A, Rehman, M Z, Umar, W, Farooqi, Z U R, Sarfraz, M, Ahmad, H R, and Aslam, M Z (2022) Role of glycine betaine in stress management in plants. In Emerging Plant Growth Regulators in Agriculture, pp. 335-356. Academic Press.

Ayub, Q, Mehmood, A, Hayat, U, Shahzad, Q, and Ahmad, S (2020) Effect of salinity on physiological and biochemical attributes of different Brinjal (Solanum melongena L.) cultivars. Pure and Applied Biology (PAB), 9(4):2190-2198.

Ayuso-Calles M, Flores-Félix JD, Amaro F, García-Estévez I, Jiménez-Gómez A, de Pinho PG, et al. (2023) Effect of Rhizobium mechanisms in improving tolerance to saline stress in lettuce plants. Chemical and Biological Technologies in Agriculture 10(1):89.

Azameti, M K, Tanuja, N, Kumar, S, Rathinam, M, Imoro, A W M, Singh, P K, and Padaria, J C (2024) Transgenic tobacco plants overexpressing a wheat salt stress root protein (TaSSRP) exhibit enhanced tolerance to heat stress. Molecular Biology Reports, 51(1):791.

Azeem, M, Shoujun, Y, Qasim, M, Abbasi, M W, Ahmed, N, Hanif, T, and Dong, R (2021) Foliar enrichment of potassium and boron overcomes salinity barriers to improve growth and yield potential of cotton (Gossypium hirsutum L.). Journal of Plant Nutrition, 44(3):438-454.

Balasubramaniam, T, Shen, G, Esmaeili, N, and Zhang, H (2023) Plants’ response mechanisms to salinity stress. Plants, 12(12):2253.

Banyal, R, Sanwal, S K, Sharma, P C, Yadav, R K, and Dagar, J C (2019) Fruit and vegetable-based saline agricultural systems for nutritional and livelihood security. In Research Developments in Saline Agriculture, pp. 729-751.

Bayoumi, Y, Abd-Alkarim, E, El-Ramady, H, El-Aidy, F, Hamed, E S, Taha, N, and Rakha, M (2021) Grafting improves fruit yield of cucumber plants grown under combined heat and soil salinity stresses. Horticulturae, 7(3):61.

Begum, N, Qin, C, Ahanger, M A, Raza, S, Khan, M I, Ashraf, M, and Zhang, L (2019) Role of arbuscular mycorrhizal fungi in plant growth regulation: implications in abiotic stress tolerance. Frontiers in Plant Science, 10:1068.

Behera, T K, Krishna, R, Ansari, W A, Aamir, M, Kumar, P, Kashyap, S P, and Kole, C (2022) Approaches involved in the vegetable crops salt stress tolerance improvement: Present status and way ahead. Frontiers in Plant Science, 12:787292.

Ben Gaied, R, Brígido, C, Sbissi, I, and Tarhouni, M (2024) Sustainable strategy to boost legumes growth under salinity and drought stress in semi-arid and arid regions. Soil Systems, 8(3):84.

Bernstein L (1959) Salt tolerance of vegetable crops.

Bernstein L, Ayers AD (1953) Salt tolerance of five varieties of onions.

Bernstein, L (1959) Salt tolerance of vegetable crops in the West. USDA Information Bulletin, 205.

Bhati, S, Chaudhary, S, and Garg, G (2021) Effect of soil salinity on growth parameters and antioxidant activity in two genotypes of eggplant (Solanum melongena L.). International Journal of Agricultural and Applied Sciences, 2:95-102.

Bijalwan, P (2021) Plant tissue culture-A new tool for vegetable improvement (Indian scenario): A review. Agricultural Reviews, 42:235-239.

Birhanu, M W (2020) Transgenic Approaches of Improving Tomato (Solanum lycopersicum) to Salt Stress Tolerance. J. Biol. Agric. Healthc., 10:1-10.

Bogoutdinova, L R, Khaliluev, M R, Chaban, I A, Gulevich, A A, Shelepova, O V, and Baranova, E N (2024) Salt Tolerance Assessment of Different Tomato Varieties at the Seedling Stage. Horticulturae, 10(6):598.

Bogoutdinova, L R, Lazareva, E M, Chaban, I A, Kononenko, N V, Dilovarova, T, Khaliluev, M R, and Baranova, E N (2020) Salt stress-induced structural changes are mitigated in transgenic tomato plants over-expressing superoxide dismutase. Biology, 9(9):297.

Bolaños, L, Martín, M, El-Hamdaoui, A, Rivilla, R, and Bonilla, I (2006) Nitrogenase inhibition in nodules from pea plants grown under salt stress occurs at the physiological level and can be alleviated by B and Ca. Plant and Soil, 280:135-142.

Bonilla, I, El-Hamdaoui, A, and Bolaños, L (2004) Boron and calcium increase Pisum sativum seed germination and seedling development under salt stress. Plant Soil, 267:97-107.

Boorboori, M R, and Li, J (2025) The effect of salinity stress on tomato defense mechanisms and exogenous application of salicylic acid, abscisic acid, and melatonin to reduce salinity stress. Soil Science and Plant Nutrition, 71(1):93-110.

Borsani, O, Cuartero, J, Victoriano, V, and M A Botella (2002) Tomato tos1 mutation identifies a gene essential for osmotic tolerance and abscisic acid sensitivity. The Plant Journal, 32:905-914.

Borucki, W, and Sujkowska, M (2008) The effects of sodium chloride-salinity upon growth, nodulation, and root nodule structure of pea (Pisum sativum L.) plants. Acta Physiol. Plant., 30:293-301.

Bose C, Das PK, Roylawar P, Rupawate P, Khandagale K, Nanda S, Gawande S (2025) Identification and analysis of the GATA gene family in onion (Allium cepa L.) in response to chromium and salt stress. BMC Genomics 26(1):1-15.

Bouzroud, S, Henkrar, F, Fahr, M, and Smouni, A (2023) Salt stress responses and alleviation strategies in legumes: a review of the current knowledge. 3 Biotech, 13(8):287.

Butt, M, Sattar, A, Abbas, T, Hussain, R, Ijaz, M, Sher, A, and Zuan, A T K (2021) Morpho-physiological and biochemical attributes of Chili (Capsicum annum L.) genotypes grown under varying salinity levels. PLoS One, 16(11):e0257893.

Büyükdinç, D T, Kantoğlu, K Y, Kuşvuran, Ş, İpek, A, Karataş, A, and Ellialtıoğlu, Ş Ş (2022) Selection of salt tolerant lines at cell level using gamma ray with callus and suspension culture techniques in black carrots (Daucus carota L. ssp. sativus var. atrorubens Alef.). Applied Radiation and Isotopes, 190:110523.

Cerruti, E, Gisbert, C, Drost, H G, Valentino, D, Portis, E, Barchi, L, and Catoni, M (2021) Grafting vigour is associated with DNA de-methylation in eggplant. Horticulture Research, 8.

Chauhan, A, Sharma, D, and Banyal, S K (2021) Potential, Challenges and Strategies Involved in Gene Introgression from Wild Relatives of Vegetable Crops: A Review. Agricultural Reviews, 42:390-397.

Chen, Z, Li, X, Zhou, R, Hu, E, Peng, X, Jiang, F, and Wu, Z (2024) An Optimized Protocol for Comprehensive Evaluations of Salt Tolerance in Crop Germplasm Accessions: A Case Study of Tomato (Solanum lycopersicum L.). Agronomy, 14(4):842.

Chourasia, K N, Lal, M K, Tiwari, R K, Dev, D, Kardile, H B, Patil, V U, and Pramanik, D (2021) Salinity stress in potato: Understanding physiological, biochemical and molecular responses. Life, 11(6):545.

Cordovilla, M P, Ligero, F, and Lluch, C (1995) Influence of host genotypes on growth, symbiotic performance and nitrogen assimilation in faba bean (Vicia faba L.) under salt stress. Plant Soil, 172:289-297.

Damasceno, L F, Cova, A M W, Gheyi, H R, ALMEIDA, W F D, Dias, J A A L, and Ribeiro, V D S (2022) Production and water consumption of eggplant under salt stress and continuous drip and pulse drip irrigation. Revista Caatinga, 35:450-459.

David E M, Lesharadevi K, Dharanidharan H, Akash S, Prabhakaran S, Firdous S, and Parthasarathi T (2022) Inducing inherent salinity tolerance of eggplants by salt induction response. F1000Research, 11 pp 244

De Abreu Araújo B, e Silva L M A, Canuto K M, Alves Filho E G, Pimentel K S, Carvalho R D C S, and Bezerra M A (2025) NMR-based metabolomics and pathway analysis of leaves and fruits of Cucumis melo L. cultivated under saline stress conditions. Food Research International, 115921

Deepa Devi N, and Arumugam T (2019) Salinity tolerance in vegetable crops: A review. Journal of Pharmacognosy and Phytochemistry, 8(3), 2717-2721

Diouf I A, Derivot L, Bitton F, Pascual L, and Causse M (2018) Water deficit and salinity stress reveal many specific QTL for plant growth and fruit quality traits in tomato. Frontiers in Plant Science, 9, 279

Dkhil B B and Denden M (2010) Salt stress induced changes in germination, sugars, starch and enzyme of carbohydrate metabolism in Abelmoschus esculentus L. (Moench.) seeds. Afr. J. Agric. Res., 5 pp 1412–1418

Dogimont C, and Sari N (2022) Special Edition-2022 Gene List for Melon. Cucurbit Genetics Cooperative Report, (45), 1-47

Duborská E, Šebesta M, Matulová M, Zvěřina O, Urík M (2022) Current strategies for selenium and iodine biofortification in crop plants. Nutrients 14(22):4717.

Dutta S, Bhattacharjya D, Sinha S, and Mandal A K (2021) Salt-Tolerant and Plant Growth-Promoting Rhizobacteria: A New-Fangled Approach for Improving Crop Yield. Harsh Environment and Plant Resilience: Molecular and Functional Aspects, 367-385

Ehtaiwwesh AF, Emsahel MJ (2020) Impact of salinity stress on germination and growth of pea (Pisum sativum L.) plants. Al-Mukhtar Journal of Science 35:146-159.

El-Aidy F, Hassan N A, El-Waraky Y, Abu El-Ftooh F, Bayoumi Y, and Elhawat N (2021) Boron, manganese and zinc reduce the hazardous impact of sodic-saline soil on growth and yield of pea (Pisum sativum L.). Journal of Plant Nutrition, 44(16), 2447-2463

El-Beltagi H S, El-Nady M F, Al-Daej M I, El-Naqma K A, Rezk A A, El-Afry M M, and Metwaly M M S (2024) Exogenous application of manganese and arginine alleviates the adverse effects of salinity stress on pea (Pisum sativum L.). Cogent Food and Agriculture, 10(1), 2389445

El-Hamdaoui A, Redondo-Nieto M, Torralba B, Rivilla R, Bonilla I, and Bolaños L (2003) Influence of boron and calcium on the tolerance to salinity of nitrogen fixing pea plants. Plant Soil, 251 pp 93–103

El-Nakhel C, Cozzolino E, Ottaiano L, Petropoulos S A, Nocerino S, Pelosi M E, and Di Mola I (2022) Effect of biostimulant application on plant growth, chlorophylls and hydrophilic antioxidant activity of spinach (Spinacia oleracea L.) grown under saline stress. Horticulturae, 8 pp 971

El-Ramady H, Prokisch J, Mansour H, Bayoumi YA, Shalaby TA, Veres S, Brevik EC (2024) Review of crop response to soil salinity stress: Possible approaches from leaching to nano-management. Soil Systems 8(1):11.

Elwan M W M (2010) Ameliorative effects of di-potassium hydrogen orthophosphate on salt-stressed eggplant. J. Plant Nutr., 33 pp 1593–1604

Eraslan F, Inal A, Pilbeam DJ, Gunes A (2008) Interactive effects of salicylic acid and silicon on oxidative damage and antioxidant activity in spinach (Spinacia oleracea L. cv. Matador) grown under boron toxicity and salinity. Plant Growth Regulation 55:207-219.

Fareed S, Haider A, Ramzan T, Ahmad M, Younis A, Zulfiqar U, and Soufan W (2024) Investigating the growth promotion potential of biochar on pea (Pisum sativum) plants under saline conditions. Scientific Reports, 14(1), 10870

Farhangi-Abriz S, and Ghassemi-Golezani K (2019) Jasmonates: mechanisms and functions in abiotic stress tolerance of plants. Biocatalysis and Agricultural Biotechnology, 20 pp 101-210

Flowers T J (2004) Improving crop salt tolerance. Journal of Experimental Botany, 55(396), 307-319

Foolad M R and Lin G Y (1997) Absence of a genetic relationship between salt tolerance during seed germination and vegetative growth in tomato. Plant Breeding, 116 pp 363-367

Foolad M R, Lin G Y, and Chen F Q (1999) Comparison of QTLs for seed germination under non-stress, cold stress and salt stress in tomato. Plant Breeding, 118 pp 167-173

Fredrickson D L and Epstein E (1975) Genetically controlled differences in salt tolerance of two species of the tomato. Plant Physiol., 56 (Suppl. 2), 4

Fu Q, Liu C, Ding N, Lin Y, and Guo B (2010) Ameliorative effects of inoculation with the plant growth-promoting rhizobacterium Pseudomonas sp. DW1 on growth of eggplant (Solanum melongena L.) seedlings under salt stress. Agric. Water Manag., 97 pp 1994–2000

García-Martí M, Piñero M C, García-Sanchez F, Mestre T C, López-Delacalle M, Martínez V, and Rivero R M (2019) Amelioration of the oxidative stress generated by simple or combined abiotic stress through the K+ and Ca2+ supplementation in tomato plants. Antioxidants, 8(4), 81

Gavrilescu M (2021) Water, soil, and plants interactions in a threatened environment. Water, 13(19), 2746

Gil-Muñoz F, Pérez-Pérez J G, Quinones A, Primo-Capella A, Cebolla J, Forner-Giner M Á, and del Mar Naval M (2020) A cross population between D. kaki and D. virginiana shows high variability for saline tolerance and improved salt stress tolerance. PLOS One, 15(2), e0229023

Giordano M, Petropoulos S A, and Rouphael Y (2021) Response and defence mechanisms of vegetable crops against drought, heat and salinity stress. Agriculture, 11(5), 463

Gowayed MH, Al-Zahrani HS, Metwali EM (2017) Improving the salinity tolerance in potato (Solanum tuberosum) by exogenous application of silicon dioxide nanoparticles. International Journal of Agriculture and Biology 19(1).

Gruda N S, Dong J, and Li X (2024) From salinity to nutrient-rich vegetables: strategies for quality enhancement in protected cultivation. Critical Reviews in Plant Sciences, 43(5), 327-347

Guo M, Wang X S, Guo H D, Bai S Y, Khan A, Wang X M, and Li J S (2022) Tomato salt tolerance mechanisms and their potential applications for fighting salinity: A review. Frontiers in Plant Science, 13, 949541

Habib SH, Kausar H, Saud HM (2016) Plant growth‐promoting rhizobacteria enhance salinity stress tolerance in okra through ROS‐scavenging enzymes. BioMed Research International 2016(1):6284547.

Hamouda M M, Badr A, Ali S S, Adham A M, Ahmed H I S, and Saad-Allah K M (2023) Growth, physiological, and molecular responses of three phaeophyte extracts on salt-stressed pea (Pisum sativum L.) seedlings. Journal of Genetic Engineering and Biotechnology, 21(1), 32

Hannachi S, and Van Labeke M C (2018) Salt stress affects germination, seedling growth and physiological responses differentially in eggplant cultivars (Solanum melongena L.). Scientia Horticulturae, 228, 56-65

Hao S, Wang Y, Yan Y, Liu Y, Wang J, and Chen S (2021) A review on plant responses to salt stress and their mechanisms of salt resistance. Horticulturae, 7(6), 132

Haq IU, Azam N, Ashraf M, Javaid MM, Murtaza G, Ahmed Z, et al. (2023) Improving the genetic potential of okra (Abelmoschus esculentus L.) germplasm to tolerate salinity stress. Scientific Reports 13(1):21504.

Hasanuzzaman M, Bhuyan M B, Zulfiqar F, Raza A, Mohsin S M, Mahmud J A, and Fotopoulos V (2020) Reactive oxygen species and antioxidant defense in plants under abiotic stress: Revisiting the crucial role of a universal defense regulator. Antioxidants, 9 pp 681

Hassan M N, Mekkawy S A, Mahdy M, Salem K F, and Tawfik E (2021) Recent molecular and breeding strategies in lettuce (Lactuca spp.). Genetic Resources and Crop Evolution, 68, 3055-3079

Hassan MU, Nawaz M, Barbanti L, Masood S (2025) Enhancing salinity tolerance in crop plants through agronomic, genetic, molecular, and physiological approaches. Frontiers in Plant Science 16:1554509.

He Y, Zhu Z, Yang J, Ni X, and Zhu B (2009) Grafting increases the salt tolerance of tomato by improvement of photosynthesis and enhancement of antioxidant enzymes activity. Environ. Exp. Bot., 66 pp 270–278

Hegazi AM, El-Shraiy AM (2017) Stimulation of photosynthetic pigments, anthocyanin, antioxidant enzymes in salt-stressed red cabbage plants by ascorbic acid and potassium silicate. Middle East Journal of Agriculture Research 6(2):553-568.

Huang Y, Zhu J, Zhen A, Chen L, and Bie Z L (2009) Organic and inorganic solutes accumulation in the leaves and roots of grafted and ungrafted cucumber plants in response to NaCl stress. J. Food Agric. Environ., 7 pp 703–708

Huma T, Shaheen T, Asghar A, Nurjis F, Khan A S, and Mukhtar Z (2015) Expression of synthetic hsr1 gene in transgenic tobacco (Nicotiana tabacum) for enhanced tolerance to drought and salt stresses. Int J Agricult Biol, 17(05), 1031-1036

Ikram M, Minhas A, Al-Huqail A A, Ghoneim A M, Mahmood S, Mahmoud E, and Ali W (2024) Promoting tomato resilience: effects of ascorbic acid and sulfur-treated biochar in saline and non-saline cultivation environments. BMC Plant Biology, 24(1), 1053

Imran S, Sarker P, Mahamud M A, Paul N C, Chakrobortty J, Harine I J, and Rahimi M (2024) Copper mitigates salinity stress by regulating water status, photosynthetic pigments and ion homeostasis and increases the yield of eggplant (Solanum melongena). BMC Plant Biology, 24(1), 927

Jaarsma R, De Boer AH (2018) Salinity tolerance of two potato cultivars (Solanum tuberosum) correlates with differences in vacuolar transport activity. Frontiers in Plant Science 9:737.

Jameel J, Anwar T, Majeed S, Qureshi H, Siddiqi E H, Sana S, and Ali H M (2024) Effect of salinity on growth and biochemical responses of brinjal varieties: implications for salt tolerance and antioxidant mechanisms. BMC Plant Biology, 24(1), 128

Jeon D, Kim J B, Kang B C, and Kim C (2023) Deciphering the genetic mechanisms of salt tolerance in Sorghum bicolor L.: key genes and SNP associations from comparative transcriptomic analyses. Plants, 12(14), 2639

Johnson R, and Puthur J T (2021) Seed priming as a cost effective technique for developing plants with cross tolerance to salinity stress. Plant physiology and biochemistry, 162, 247-257

Kabała K, Reda M, Wdowikowska A, and Janicka M (2022) Role of plasma membrane NADPH oxidase in response to salt stress in cucumber seedlings. Antioxidants, 11(8), 1534

Kadoglidou K, Xanthopoulou A, Kalyvas A, and Mellidou I (2021) Utilization of tomato landraces to improve seedling performance under salt stress. Stresses, 1 pp 238-252

Kamińska I, Lukasiewicz A, Klimek-Chodacka M, Długosz-Grochowska O, Rutkowska J, Szymonik K, Baranski R (2022) Antioxidative and osmoprotecting mechanisms in carrot plants tolerant to soil salinity. Scientific Reports 12(1):7266.

Kapazoglou A, Gerakari M, Lazaridi E, Kleftogianni K, Sarri E, Tani E, and Bebeli P J (2023) Crop wild relatives: A valuable source of tolerance to various abiotic stresses. Plants, 12 pp 328

Kappel N, Boros I F, Ravelombola F S, and Sipos L (2021) EC sensitivity of hydroponically-grown lettuce (Lactuca sativa L.) types in terms of nitrate accumulation. Agriculture, 11 pp 315

Kesawat M S, Satheesh N, Kherawat B S, Kumar A, Kim H U, Chung S M, and Kumar M (2023) Regulation of reactive oxygen species during salt stress in plants and their crosstalk with other signaling molecules—Current perspectives and future directions. Plants, 12(4), 864

Khalid M F, Huda S, Yong M, Li L, Li L, Chen Z H, and Ahmed T (2023) Alleviation of drought and salt stress in vegetables: crop responses and mitigation strategies. Plant Growth Regulation, 99 pp 177-194

Khan M A H, Baset Mia M A, Quddus M A, Sarker K K, Rahman M, Skalicky M, and Hossain A (2022) Salinity-induced physiological changes in pea (Pisum sativum L.): Germination rate, biomass accumulation, relative water content, seedling vigor and salt tolerance index. Plants, 11(24), 3493

Khan P S V, and Basha P O (2015) Salt stress and leguminous crops: Present status and prospects. Legumes under Environmental Stress: Yield, Improvement and Adaptations, 21-51

Kumar P, and Sharma P K (2020) Soil salinity and food security in India. Frontiers in sustainable food systems, 4, 533781

Kumar S, Li G, Yang J, Huang X, Ji Q, Liu Z, and Hou H (2021) Effect of salt stress on growth, physiological parameters, and ionic concentration of water dropwort (Oenanthe javanica) cultivars. Frontiers in plant science, 12, 660409

Kumari P V, and Mesfin Y (2015) Testing salt tolerance to boost on chickpea (Cicer arietinum L. Mill Sp) biomass/cultivation. Global Journal of Research on Medicinal Plants and Indigenous Medicine, 4 pp 79

Kumawat K C, Sharma B, Nagpal S, Kumar A, Tiwari S, and Nair R M (2023) Plant growth-promoting rhizobacteria: Salt stress alleviators to improve crop productivity for sustainable agriculture development. Frontiers in Plant Science, 13 pp 1101862

Lahlali R, Laasli S E, and Ait Barka E (2025) Plant responses to biotic and abiotic stresses: From cellular to morphological changes—Series II. Agronomy, 15(1), 229

Levy D (1992) The response of potatoes (Solanum tuberosum L.) to salinity: plant growth and tuber yields in the arid desert of Israel. Annals of Applied Biology 120(3):547-555.

Li N, Cui Y, Zhang Z, Wang S, Sun Y, Zhang S, and Li G (2024) Overexpression of vacuolar H+-pyrophosphatase from a recretohalophyte Reaumuria trigyna enhances vegetative growth and salt tolerance in transgenic Arabidopsis thaliana. Frontiers in Plant Science, 15, 1435799

Li W Q, Li J Y, Bi S J, Jin J Y, Fan Z L, Shang Z L, and Wang Y J (2025) Melatonin enhances maize germination, growth, and salt tolerance by regulating reactive oxygen species accumulation and antioxidant systems. Plants, 14(2), 296

Li Y, Wu X, Zhang Y, Zhang Q (2022) CRISPR/Cas genome editing improves abiotic and biotic stress tolerance of crops. Frontiers in Genome Editing 4:987817.

Lin C, Zheng S, Liu K, Yu R, Guan P, Hu B, and Zhang X (2025) Elucidating the molecular basis of salt tolerance in potatoes through miRNA expression and phenotypic analysis. Scientific Reports, 15(1), 2635

Lin S, Yang J, Liu Y, and Zhang W (2024) MsSPL12 is a positive regulator in alfalfa (Medicago sativa L.) salt tolerance. Plant Cell Reports, 43(4), 101

Liu C, Mao B, Yuan D, Chu C, and Duan M (2022) Salt tolerance in rice: Physiological responses and molecular mechanisms. The Crop Journal, 10 pp 13-25

Liu H, Sun J, Zou J, Li B, and Jin H (2023) MeJA-mediated enhancement of salt-tolerance of Populus wutunensis by 5-aminolevulinic acid. BMC Plant Biology, 23(1), 185

Mady E, Abd El-Wahed A H, Awad A H, Asar T O, Al-Farga A, Abd El-Raouf H S, and Hamada F A (2023) Evaluation of salicylic acid effects on growth, biochemical, yield, and anatomical characteristics of eggplant (Solanum melongena L.) plants under salt stress conditions. Agronomy, 13(9), 2213

Malcolm CV, Smith ST (1971) Growing plants with salty water. Journal of the Department of Agriculture, Western Australia, Series 4 12(2):41-44.

Martínez J P, Fuentes R, Badilla D, Rosales C, Alfaro-Quezada J F, Correa F, and Lutts S (2024) Improving the salt tolerance of “Old limachino tomato” by using a new salt-tolerant rootstock. Horticulturae, 10(8), 780

Massa R, Al-Biski F, Al-Maarri K (2024) Clonal selection for salinity tolerance in some potato (Solanum tuberosum) varieties by callus culture. Potato Research 67(2):647-661.

Matera A, Warchoł M, Simlat M (2025) Effect of the Pantoea vagans strain SRS89 on carrot (Daucus carota subsp. sativus L.) seed germination and plant growth under saline conditions. South African Journal of Botany 180:415-427.

Mendonça A J, Lima G S D, Soares L A D A, Oliveira V K, Gheyi H R, Silva L D A, and Fernandes P D (2023) Gas exchange, photosynthetic pigments, and growth of hydroponic okra under salt stress and salicylic acid. Revista Brasileira de Engenharia Agrícola e Ambiental, 27, 673-681

Mohammed D, Maina MM, Audu I, Tudunwada IY, Nasiru NK, Nasidi NM, Umar SE (2023) Application of multitemporal Landsat data in mapping of saline soil in Kano River Irrigation Scheme (KRIS). FUDMA Journal of Sciences 7(1):193-200.

Mondal S, and Chakraborty K (2020) Brassicaceae plants response and tolerance to salinity. The Plant Family Brassicaceae: Biology and Physiological Responses to Environmental Stresses, 203-228

Mondal, S, Rahaman, E H M S and Asch, F (2022). Potassium content is the main driver for salinity tolerance in sweet potato before tuber formation. Journal of Agronomy and Crop Science, 208(5), 645-661.

Moreels, P, Bigot, S, Defalque, C, Correa, F, Martinez, J P, Lutts, S and Quinet, M (2023). Intra-and inter-specific reproductive barriers in the tomato clade. Frontiers in Plant Science, 14, 1326689.

Motamedi, M, Azizpour, K and Sarabi, V (2021). The effects of salinity stress caused by different levels of NaCl on okra. Genetic Engineering and Biosafety Journal, 10, 68-81.

Muhammad, M, Waheed, A, Wahab, A, Majeed, M, Nazim, M, Liu, Y H, ... and Li, W J (2024). Soil salinity and drought tolerance: An evaluation of plant growth, productivity, microbial diversity, and amelioration strategies. Plant Stress, 11, 100319.

Mushtaq, Z, Faizan, S and Gulzar, B (2020). Salt stress, its impacts on plants and the strategies plants are employing against it: A review. Journal of Applied Biology and Biotechnology, 8, 81-91.

Nadler A, Heuer B (1995) Effect of saline irrigation and water deficit on tuber quality. Potato Research 38:119-123.

Naeem, M, Iqbal, M, Shakeel, A, Ul-Allah, S, Hussain, M, Rehman, A and Ashraf, M (2020). Genetic basis of ion exclusion in salinity stressed wheat: Implications in improving crop yield. Plant Growth Regulation, 92, 479-496.

Nagaz K, Masmoudi MM, Mechlia NB (2012) Impacts of irrigation regimes with saline water on carrot productivity and soil salinity. Journal of the Saudi Society for Agricultural Sciences 11(1):19-27.

Najafi, R, Rezaei, A and Mozafarian, M (2024). Physiological and biochemical responses of okra (Abelmoschus esculentus) under salinity stress in Iran. Journal of Agriculture and Food Research, 18, 101322.

Neto HDSL, Garita SA, Bernardo VF, Ruscitti MF, Arango MC, Wahnan LK, de Almeida Guimarães M (2024) Salicylic and jasmonic acid attenuate the salt stress effects in lettuce.

Nusrat, N, Shahbaz, M and Perveen, S (2014). Modulation in growth, photosynthetic efficiency, activity of antioxidants, and mineral ions by foliar application of glycinebetaine on pea (Pisum sativum L.) under salt stress. Acta physiologiae plantarum, 36, 2985-2998.

Omar MM, Taha AA, El-Eraky SMR (2020) Combined effect of some nitrogen and zinc sources on some quality parameters of spinach grown on salt-affected soil. Journal of Soil Sciences and Agricultural Engineering 11(6):215-221.

Orosco-Alcalá, B E, Núñez-Palenius, H G, Díaz-Serrano, F, Pérez-Moreno, L, Valencia-Posadas, M, Trejo-Tellez, L I and Valiente-Banuet, J I (2021). Grafting improves salinity tolerance of bell pepper plants during greenhouse production.