Benefits of natural biostimulants: Ginger and moringa extracts as seed priming agents to improve salinity tolerance of Ocimum basilicum L.
DOI:
https://doi.org/10.15835/nbha53414869Keywords:
biostimulant, ginger extract, moringa extract, Ocimum basilicum L., priming, salinity, secondary metabolismAbstract
Using natural biostimulants such as plant extracts is a novel agronomic biofortification strategy to mitigate abiotic stresses such as salinity on crops. This study assessed seed priming effects on physiological and biochemical attributes of basil (Ocimum basilicum L.) under salinity (0 and 100 mM NaCl). Seeds were primed for 12h with distilled water (hydro-priming, H2O), moringa leaf (ME) and ginger rhizome extracts (GE). Salt stress reduced plant fresh weight and hydration, but priming with all agents alleviated salinity impact on basil. ME priming notably boosted fresh weight (+81%) and water content (+50%) compared to unprimed salt-exposed plants. ME-primed plants exhibited the highest proline (2-fold) and sugar (1.5-fold) contents contributing to the osmotic adjustment. Seed priming with ME also reduced malondialdehyde level (MDA) by about 50% enhancing membrane stability. ME also significantly modulated secondary metabolism through the stimulation of polyphenols (+88%), flavonoids (+98%) and total antioxidant activity (+1.4-fold). It upregulated key enzymes: Phenylalanine ammonia-lyase (PAL, +77%), Shikimate dehydrogenase (SKDH, +119%) and Polyphenoloxidase (PPO, +133%). Correlation analysis indicated that priming generally shifted correlation values from negative under salt stress to positive, with most r coefficients ranging between 0.8 and 1.00. Principal Component Analysis (PCA) clearly separated the primed and unprimed conditions, with the first two components explaining over 90% of the variability. Hence, these multivariate approaches confirm the beneficial effect of seed priming on basil growth and metabolism under salinity. Taken together, our findings provide new insights into the potential of using moringa leaf extract as a biostimulant to alleviate environmental stresses.
References
Abd El-Azim WM, Khater RMR, Badawy MYM (2017). Effect of bio-fertilization and different licorice extracts on growth and productivity of Foeniculum vulgare, Mill. Plant. Middle East Journal of Agriculture Research 6:1-12. https://www.curresweb.com/mejar/mejar/2017/1-12.pdf
Ahmed T, Elezz AA, Khalid MF (2021). Hydropriming with moringa leaf extract mitigates salt stress in wheat seedlings. Agriculture 11(12):1254. https://doi.org/10.3390/agriculture11121254
Al Khazan MM (2020). Priming with moringa (Moringa oleifera lam.) leaf extract boosts the growth and physio-biochemical attributes of lead-stressed fenugreek (Trigonella foenum-graecum L.) seedlings. Applied Ecology and Environmental Research 18(5):6949-6967. https://doi.org/10.15666/aeer/1805_69496967
Al Otaibi F, Alghamdi S, Abo-Elyousr K (2024). The influence of salinity on plant growth and amendment strategies. Sohag Journal of Sciences 9(3):261-267. https://doi.org/10.21608/sjsci.2024.258471.1168
Albaladejo-Marico L, Carvajal M, Yepes-Molina L (2025). Priming with broccoli extract mitigates salinity stress in tomato through enhanced water and Na+ / K+ homeostasis. Journal of Plant Physiology 314:154612. https://doi.org/10.1016/j.jplph.2025.154612
Ali EF, Al-Yasi HM, Issa AA, Hessini K, Hassan FAS (2022). Ginger extract and fulvic acid foliar applications as novel practical approaches to improve the growth and productivity of Damask Rose. Plants 11(3):412. https://doi.org/10.3390/plants11030412
Arraf E, Al-madhagi I, Haissam H (2025). Plant extracts are a promising solution to alleviate salinity stress at the Chili pepper germination stage. Sana’a University Journal of Applied Sciences and Technology 3(3):868-884. https://doi.org/10.59628/jast.v3i3.1663
Bakhoum G, Sadak M, Tawfic M (2022). Chitosan and chitosan nanoparticle effect on growth, productivity and some biochemical aspects of Lupinus termis L. plant under drought conditions. Egyptian Journal of Chemistry 65(5):537-549. https://doi.org/10.21608/ejchem.2021.97832.4563
Bates LS, Wadern RP, Teare ID (1973). Rapid determination of free proline for water stress studies. Plant and Soil 39:205-207. https://doi.org/10.1007/BF00018060
Bounaouara F, Hidri R, Abdallah M, Falouti M, Rabhi M, Selmi S, … Zorrig W (2025). The Impact of exogenous silicon on phenolic compound metabolism in Sorghum: a qualitative and quantitative analysis. Journal of Soil Science and Plant Nutrition 25:5508-5532 https://doi.org/10.1007/s42729-025-02474-0
Bradford MM (1976). A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry 72(1-2):248-254. https://doi.org/10.1016/0003-2697(76)90527-3
Chakhchar A, Lamaoui M, Wahbi S, Ferradous A, El Mousadik A, Ibnsouda-Koraichi S, … El Modafar C (2015). Leaf water status, osmoregulation and secondary metabolism as a model for depicting drought tolerance in Argania spinosa. Acta Physiologiae Plantarum 37:1-16. https://doi.org/10.1007/s11738-015-1833-8
Cheng F, Cheng Z (2015). Research progress on the use of plant allelopathy in agriculture and the physiological and ecological mechanisms of allelopathy. Frontiers in Plant Science 6:1020. https://doi.org/10.3389/fpls.2015.01020
Di Sario L, Boeri P, Matus JT (2025). Plant biostimulants to enhance abiotic stress resilience in crops. International Journal of Molecular Sciences 26(3):1129. https://doi.org/10.3390/ijms26031129
Ellouzi H, Oueslati S, Hessini K, Rabhi M, Abdelly C (2021). Seed-priming with H2O2 alleviates subsequent salt stress by preventing ROS production and amplifying antioxidant defense in cauliflower seeds and seedlings. Scientia Horticulturae 288:110360. https://doi.org/10.1016/j.scienta.2021.110360
Ellouzi H, Rabhi M, Khedher S, Debez A, Abdelly C. Zorrig W (2023). Silicon seed priming enhances salt tolerance of barley seedlings through early ROS detoxification and stimulation of antioxidant defence. Silicon 15(1):37-60. https://doi.org/10.1007/s12633-022-02001-1
FAO Food and Agriculture Organization of the United Nations (2009). How to feed the world 2050. In: Proceedings of the Expert Meeting on How to Feed the World in 2050. FAO Headquarters, Rome https://www.fao.org/4/ak542e/ak542e00.pdf
Gassoumi W, Ellouzi H, Ben Slimene I, Zar Kalai F, Ben Ayed R, Zorrig W, … Oueslati S (2024). Plant extracts and plant growth promoting rhizobacteria : seed bio‑priming approach to improve tolerance of Lactuca sativa L. under salt stress. Euro-Mediterranean Journal for Environmental Integration 10:1901-1918. https://doi.org/10.1007/s41207-024-00633-w
Gedeon S, Ioannou A, Balestrini R, Fotopoulos V, Chrystalla A (2022). Application of biostimulants in tomato plants (Solanum lycopersicum) to enhance plant growth and salt stress tolerance. Plants 11(22):3082. https://doi.org/10.3390/plants11223082
Ghezal N, Rinez I, Sbai H, Saad I, Farooq M, Rinez A, … Haouala R (2016). Improvement of Pisum sativum salt stress tolerance by bio-priming their seeds using Typha angustifolia leaves aqueous extract. South African Journal of Botany 105:240-250. https://doi.org/10.1016/j.sajb.2016.04.006
Guo X, Ahmad N, Zhao S, Zhao C, Zhong W, Wang X, Li G (2022). Effect of salt stress on growth and physiological properties of Asparagus seedlings. Plants 11(21):1-14. https://doi.org/10.3390/plants11212836
Hasanuzzaman M, Bhuyan MHMB, Zulfiqar F, Raza A, Mohsin SM, Al Mahmud J, … 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(8):1-52. https://doi.org/10.3390/antiox9080681
Hasanuzzaman M, Parvin K, Bardhan K, Nahar K, Anee TI (2021). Biostimulants for the regulation of reactive oxygen species metabolism in plants under abiotic stress. Cells 10(10):2537. https://doi.org/10.3390/cells10102537
Hassan F, Al-yasi H, Ali E, Alamer K, Hessini K, Attia H, El-shazly S (2021). Mitigation of salt-stress effects by moringa leaf extract or salicylic acid through motivating antioxidant machinery in Damask rose. Canadian Journal of Plant Science 101: 157-165. https://doi.org/10.1139/cjps-2020-0127
Hewitt EJ (1966). Sand and water culture methods used in the study of plant nutrition. In: Experimental research. Farnham Royal, England: Commonwealth Agricultural Bureaux (2nd ed) pp. 547. https://doi.org/10.1017/s0014479700021852
Hussain S, Khan F, Cao W, Wu L, Geng M (2016). Seed priming alters the production and detoxification of reactive oxygen intermediates in rice seedlings grown under sub-optimal temperature and nutrient supply. Frontiers in Plant Science 7:439. https://doi.org/10.3389/fpls.2016.00439
Jawaid MZ, Khalid MF, Elezz AA, Ahmed T (2025). Seed priming mitigates the salt stress in eggplant (Solanum melongena) by activating antioxidative defense mechanisms. Phyton-International Journal of Experimental Botany 94(8):2423-2439. https://doi.org/10.32604/phyton.2025.068303
Kamanga BM, Cartmill DL, McGill C, McCormick AC (2025). Allelopathic effects of Moringa oleifera Lam. on cultivated and non-cultivated plants: Implications for crop productivity and sustainable agriculture. Agronomy 15(8):1766. https://doi.org/10.3390/agronomy15081766
Kamran M, Xie K, Sun J, Wang D, Shi C, Lu Y, Gu W, Xu P (2020). Modulation of growth performance and coordinated induction of ascorbate-glutathione and methylglyoxal detoxification systems by salicylic acid mitigates salt toxicity in choysum (Brassica parachinensis L.). Ecotoxicology and Environmental Safety 188:109877. https://doi.org/10.1016/j.ecoenv.2019.109877
Khalofah A, Bokhari NA, Migdadi HM, Alwahibi MS (2020). Antioxidant responses and the role of Moringa oleifera leaf extract for mitigation of cadmium stressed Lepidium sativum L. South African Journal of Botany 129:341-346. https://doi.org/10.1016/j.sajb.2019.08.041
Khan S, Basra SMA, Afzal I, Nawaz M, Rehman HU (2017). Growth promoting potential of fresh and stored Moringa oleifera leaf extracts in improving seedling vigor, growth and productivity of wheat crop. Environmental Science and Pollution Research 24(35):27601-27612. https://doi.org/10.1007/s11356-017-0336-0
Khan S, Ibrar D, Bashir S, Rashid N, Hasnain Z, Nawaz M, … Dvořáček J (2022). Application of Moringa leaf extract as a seed priming agent enhances growth and physiological attributes of Rice seedlings cultivated under water deficit regime. Plants 11(3):1-17. https://doi.org/10.3390/plants11030261
Li G, Song C, Manzoor MA, Li D, Cao Y, Cai Y (2023). Functional and kinetics of two efficient phenylalanine ammonia lyase from Pyrus bretschneideri. BMC Plant Biology 23:612. https://doi.org/10.1186/s12870-023-04586-0
Liu H, Able AJ, Able JA (2022). Priming crops for the future: rewiring stress memory. Trends in Plant Science 27(7):699-716. https://doi.org/10.1016/j.tplants.2021.11.015
Noor A, Wahocho NA, Haider N, Jamali MF, Talpur A, Abro M, Wahocho SA (2024). Priming with Moringa leaf extract improves the germination and growth of bottle gourd (Lagenaria siceraria L.). Pakistan Journal of Agriculture 1(1):1-5. https://doi.org/10.38211/PJA.2024.01.82
Noreen S, Saleem S, Iqbal U, Mahmood S, Salim Akhter M, Akbar N, … Kaushik P (2024). Moringa olifera leaf extract increases physio-biochemical properties, growth and yield of Pisum sativum grown under salinity stress. Journal of King Saud University- Science 36(2):103056. https://doi.org/10.1016/j.jksus.2023.103056
Oueslati S, Ellouzi H, Jday M, Zar Kalai F, Alyam, NM, Hanana M, … Debez A. (2025). Enhancing salt tolerance and growth in Brassica rapa var. rapa L. through vitamin seed priming : The role of ascorbic acid and riboflavin. Scientia Horticulturae 350:114354. https://doi.org/10.1016/j.scienta.2025.114354
Oueslati S, Serairi Beji R, Zar Kalai F, Soufiani M, Zorrig W, Aissam S, …El Modafar C (2023). Antioxidant potentialities and gastroprotective effect of Reichardia picroides extracts on Ethanol/HCl induced gastric ulcer rats. International Journal of Environmental Health Research 34(2):1088-1099. https://doi.org/10.1080/09603123.2023.2198760
Peek J, Christendat D (2015). The shikimate dehydrogenase family: Functional diversity within a conserved structural and mechanistic framework. Archives of Biochemistry and Biophysics 566:85-99. https://doi.org/10.1016/j.abb.2014.12.006
Prieto P, Pineda M, Aguilar M (1999). Spectrophotometric quantitation of antioxidant capacity through the formation of a phosphomolybdenum complex: Specific application to the determination of vitamin E. Analytical Biochemistry 269:337-341. https://doi.org/10.1006/abio.1999.4019
Rady MM, Desoky ESM, Elrys AS, Boghdad, MS (2019). Can licorice root extract be used as an effective natural biostimulant for salt-stressed common bean plants? South African Journal of Botany 121:294-305. https://doi.org/10.1016/j.sajb.2018.11.019
Raja V, Majeed U, Kang H, Andrabi KI, John R (2017). Abiotic stress: Interplay between ROS, hormones and MAPKs. Environmental and Experimental Botany 137:142-157. https://doi.org/10.1016/j.envexpbot.2017.02.010
Righini H, Francioso O, Di M, Prodi A, Martel A, Roberti R (2021). Tomato seed biopriming with water extracts from Anabaena minutissima, Ecklonia maxima and Jania adhaerens as a new agro-ecological option against Rhizoctonia solani. Scientia Horticulturae 281:109921. https://doi.org/10.1016/j.scienta.2021.109921
Shabana MH, Balbaa LK, Talaat IM (2017). Effect of foliar applications of Zingiber officinale extracts on Origanum majorana. Journal of Herbs, Spices and Medicinal Plants 23(2):89-97. https://doi.org/10.1080/10496475.2016.1269381
Singh P, Singh J, Ray S, Rajput RS, Vaishnav A, Singh RK, Singh HB (2020). Seed biopriming with antagonistic microbes and ascorbic acid induce resistance in tomato against Fusarium wilt. Microbiological Research 237:126482. https://doi.org/10.1016/j.micres.2020.126482
Soares TFSN, Da Silva AVC, Muniz EN (2021). Moringa leaf extract: A cost-effective and sustainable product to improve plant growth. South African Journal of Botany 141:171-176. https://doi.org/10.1016/j.sajb.2021.04.007
Song JN, Yang, JL, Dong XX, Zhang HJ, Jeong BR (2024). Supplemented biochar mitigates the ammonium toxicity in Basil (Ocimum basilicum L.) plants. Russian Journal of Plant Physiology 71:189. https://doi.org/10.1134/S1021443724605512
Suh HY, Yoo KS, Suh SG (2014). Effect of foliar application of fulvic acid on plant growth and fruit quality of tomato (Lycopersicon esculentum L.). Horticulture Environment and Biotechnology 55(6):455-461. https://doi.org/10.1007/s13580-014-0004-y
Xu Y, Bu W, Xu Y, Fei H, Zhu Y, Ahmad I, … Zhu G (2024). Effects of salt stress on physiological and agronomic traits of Rice genotypes with contrasting salt tolerance. Plants 13(8):1157. https://doi.org/10.3390/plants13081157
Yakhin OI, Lubyanov AA, Yakhin IA, Brown PH (2017). Biostimulants in plant science: A global perspective. Frontiers in Plant Science 7:2049. https://doi.org/10.3389/fpls.2016.02049
Yang Z, Zhi P, Chang C (2022). Priming seeds for the future: Plant immune memory and application in crop protection. Frontiers in Plant Science 13:961840. https://doi.org/10.3389/fpls.2022.961840
Younis A, Akhtar MS, Riaz A, Zulfiqar F, Qasim M, Farooq A, … Bhatti ZM (2018). Improved cut flower and corm production by exogenous moringa leaf extract application on gladiolus cultivars. Acta Scientiarum Polonorum Hortorum Cultus 17(4):25-28. https://doi.org/10.24326/asphc.2018.4.3
Zulfiqar F, Casadesús A, Brockman H, Munné-Bosch S (2020). An overview of plant-based natural biostimulants for sustainable horticulture with a particular focus on moringa leaf extracts. Plant Science 295:110194. https://doi.org/10.1016/j.plantsci.2019.110194
Downloads
Published
How to Cite
Issue
Section
License
Copyright (c) 2025 Samia OUESLATI, Hasna ELLOUZI, Walid ZORRIG, Ahmed DEBEZ, Chedly ABDELLY, Mohsen HANANA

This work is licensed under a Creative Commons Attribution 4.0 International License.
License:

Open Access Journal:
The journal allows the author(s) to retain publishing rights without restriction. Users are allowed to read, download, copy, distribute, print, search, or link to the full texts of the articles, or use them for any other lawful purpose, without asking prior permission from the publisher or the author.









.png)





