Overview of biostimulants in horticultural products under abiotic stress: A review

Document Type : Review Article

Authors

Department of Horticulture, Faculty of Agriculture, Ilam University, Ilam, Iran

Abstract
Global agriculture will face two major challenges in the future: population growth and climate change. One of the primary obstacles to agricultural productivity is environmental stress, with abiotic factors having a considerable negative effect on horticultural yields. As such, exploring effective strategies to mitigate plant stress is essential. Among these strategies, the use of biostimulants has emerged as an innovative and eco-friendly approach to boost growth and productivity in horticultural crops under stress conditions. Biostimulants, including seaweeds, fungi, bacteria, and amino acids, play a key role in alleviating stress effects. Their effectiveness is largely attributed to bioactive compounds such as polysaccharides, pigments, phenolics, proteins, phytohormones, and various micro- and macronutrients. Research indicates that these compounds can significantly reduce plant stress and enhance resilience. The effective use of bio-stimulants can reduce waste from fertilizers and minimize the risk of nutrient runoff that leads to environmental pollution. Economically, it enhances nutrient uptake, plant growth, and productivity under stress conditions, reduces input costs, and increases farmers' profitability. Therefore, the use of these compounds in greenhouse systems is not only environmentally significant but also economically important. This study delves into the application of biostimulants in horticulture under abiotic stress and highlights some of the key challenges associated with their broader adoption and implementation.

Keywords


  1. Abbas, S., Javed, M. T., Ali, Q., Azeem, M., and Ali, S. 2021. Nutrient deficiency stress and relation with plant growth and development. In Engineering tolerance in crop plants against abiotic stress (pp. 239-262). CRC Press.‏
  2. Abdelkader, M., Voronina, L., Shelepova, O., Puchkov, M., Loktionova, E., Zhanbyrshina, N., and  Ksenofontov, A. 2023. Monitoring role of exogenous amino acids on the proteinogenic and ionic responses of lettuce plants under salinity stress conditions. Horticulturae, 9(6), 626.‏ https://doi.org/10.3390/horticulturae9060626
  3. Abdou, M. A. H., Hassan, E. E., Ibrahim, A. M., and Abdel-Rahim, A. F. A. 2021. Effect of some biostimulating substances on zonal geranium growth and flowering under irrigation water salinity. Scientific Journal of Flowers and Ornamental Plants, 8(4), 497-512. 10.21608/SJFOP.2021.381297
  4. Ali, E.F., Hassan, F.A.S. and Elgimabi, M., 2018. Improving the growth, yield and volatile oil content of Pelargonium graveolens L. Herit by foliar application with moringa leaf extract through motivating physiological and biochemical parameters. South African Journal of Botany, 119, pp.383-389. https://doi.org/10.1016/j.sajb.2018.10.003
  5. Ali, O., Farrell, A. D., Ramsubhag, A., and Jayaraman, J. 2024. Beneficial effects of an Ascophyllum nodosum extract on tomato (Solanum lycopersicum L.) during water stress. Journal of Applied Phycology, 36(1), 385-397.‏ doi:10.1007/s10811-023-03156-z.
  6. Ali, A., Niu, G., Masabni, J., Ferrante, A., and Cocetta, G. 2024. Integrated Nutrient Management of Fruits, Vegetables, and Crops through the Use of Biostimulants, Soilless Cultivation, and Traditional and Modern Approaches—A Mini Review. Agriculture, 14(8), 1330. https://doi.org/10.3390/agriculture14081330
  7. Aytaç, Z., Gülbandılar, A. and Kürkçüoğlu, M. 2022. Humic acid improves plant yield, antimicrobial activity and essential oil composition of Oregano (Origanum vulgare L. subsp. hirtum (Link.) Ietswaart). Agronomy, 12(9), p.2086. doi.org/10.3390/agronomy12092086
  8. Balasubramaniam, T., Shen, G., Esmaeili, N., and Zhang, H. 2023. Plants’ response mechanisms to salinity stress. Plants, 12(12), 2253.‏ https://doi.org/10.3390/plants12122253
  9. Bayanati, M., Asgari Lajayer, B., Al-Tawaha, A. R., and Astatkie, T. 2023. Seed bio-priming: An emerging tool towards improved germination and agricultural sustainability. In New Frontiers in Plant-Environment Interactions: Innovative Technologies and Developments (pp. 253-265). Cham: Springer Nature Switzerland.‏ https://doi.org/10.1007/978-3-031-43729-8_9
  10. Bhatla, S.C.,and Lal, M.A. 2023. Abiotic Stress. In: Plant Physiology, Development and Metabolism. Springer, Singapore. https://doi.org/10.1007/978-981-99-5736-1_31
  11. Bhupenchandra, I., Chongtham, S. K., Devi, E. L., Choudhary, A. K., Salam, M. D., Sahoo, M. R., Bhutia, T.L., Devi, S.H, Thounaojam, A.S, Behera, C., M.N. H., Kumar, A., Dasgupta, M., Devi, Y.P., Singh, D., Bhagowati, S., Devi, C.P., Singh, H.R., and Khaba, C.I. 2022. Role of biostimulants in mitigating the effects of climate change on crop performance. Frontiers in Plant Science. 13:967665. doi: 10.3389/Fpls.2022.967665
  12. Bulgari, R., Franzoni, G., and Ferrante, A. 2019. Biostimulants application in horticultural crops under abiotic stress conditions. Agronomy, 9(6), 306.‏ https://doi.org/10.3390/Agronomy9060306
  13. Cardarelli, M., Woo, S. L., Rouphael, Y., and Colla, G. 2022. Seed treatments with microorganisms can have a biostimulant effect by influencing germination and seedling growth of crops. Plants, 11(3), 259.‏ https://doi.org/10.3390/Plants11030259
  14. Dalvand, M., Solgi, M., and Khaleghi, A. 2018. Effects of foliar application of humic acid and drought stress on growth and physiological characteristics of marigold (Taget erecta), J. Sci. Technol. Greenhouse Cul., 9(2), 67-79. DOI: 10.29252/ejgcst.9.2.67.
  15. De Clercq, P., Pauwels, E., Top, S., Steppe, K. and Van Labeke, M.C., 2023. Effect of seaweed-based biostimulants on growth and development of Hydrangea paniculata under continuous or periodic drought stress. Horticulturae, 9(4), p.509. https://doi.org/10.3390/horticulturae9040509
  16. Del Buono, D. 2021. Can biostimulants be used to mitigate the effect of anthropogenic climate change on agriculture? It is time to respond. Science of the Total Environment, 751, 141763.‏ https://doi.org/10.1016/j.scitotenv.2020.141763
  17. Demir, I., Ozden, E., Yıldırım, K. C., Sahin, O., and Van Staden, J. 2018. Priming with smoke-derived karrikinolide enhances germination and transplant quality of immature and mature pepper seed lots. South African Journal of Botany, 115, 264-268. https://doi.org/10.1016/J.sajb.2017.07.001
  18. Deolu‐Ajayi, A.O., van der Meer, I.M., Van der Werf, A. and Karlova, R. 2022. The power of seaweeds as plant biostimulants to boost crop production under abiotic stress. Plant, Cell and Environment, 45(9), pp.2537-2553. https://doi.org/10.1111/pce.14391
  19. Desoky, E. S. M., Elrys, A. S., and Rady, M. M. 2019. Integrative moringa and licorice extracts application improves Capsicum annuum fruit yield and declines its contaminant contents on a heavy metals-contaminated saline soil. Ecotoxicology and Environmental Safety, 169, 50-60. https://doi.org/10.1016/J.ecoenv.2018.10.117
  20. Di Sario, L.; Boeri, P.;Matus, J.T.; Pizzio, G.A. 2025. Plant Biostimulants to Enhance Abiotic Stress Resilience in Crops. Int. J. Mol. Sci. 26, 1129. https://doi.org/10.3390/ijms26031129
  21. Dos Santos, T. B., Ribas, A. F., de Souza, S. G. H., Budzinski, I. G. F., and Domingues, D. S. 2022. Physiological responses to drought, salinity, and heat stress in plants: a review. Stresses2(1), 113-135. https://doi.org/10.3390/stresses2010009
  22. Dubey, R., and Misra, S. 2024. Biostimulants: an eco-friendly regulator of plant stress tolerance and sustainable solution to future agriculture. Proceedings of the Indian National Science Academy, 1-8.‏ doi.org/10.1007/s43538-024-00328-4
  23. Elsayed, S.I.M., Sabra, A.S., and Omer, E.A. 2023. Role of Plant Extracts and Biostimulant in Mitigating Plant Drought and Salinity Stress. In: Hasanuzzaman, M. (eds) Climate-Resilient Agriculture, Vol 2. Springer, Cham. https://doi.org/10.1007/978-3-031-37428-9_25
  24. Eltahawy, A. M., Awad, E. S. A., Ibrahim, A. H., Merwad, A. R. M., and Desoky, E. S. M. 2022. Integrative application of heavy metal–resistant bacteria, moringa extracts, and nano-silicon improves spinach yield and declines its contaminant contents on a heavy metal–contaminated soil. Frontiers in Plant Science, 13, 1019014. doi: 10.3389/Fpls.2022.1019014
  25. Eshaghi Gorgi, O., Fallah, H., Niknejad, Y., and Barari Tari, D. 2022. Effect of Plant growth promoting rhizobacteria (PGPR) and mycorrhizal fungi inoculations on essential oil in Melissa officinalis L. under drought stress. Biologia, 77(1), 11-20.‏ https://doi.org/10.1007/s11756-021-00919-2
  26. Franzoni, G., Cocetta, G., Prinsi, B., Ferrante, A., and Espen, L. 2022. Biostimulants on crops: Their impact under abiotic stress conditions. Horticulturae, 8(3), 189.‏ https://doi.org/10.3390/horticulturae8030189
  27. García-García, A. L., García-Machado, F. J., Borges, A. A., Morales-Sierra, S., Boto, A., and Jiménez-Arias, D. 2020. Pure organic active compounds against abiotic stress: A biostimulant overview. Frontiers in Plant Science, 11, 575829.‏ doi: 10.3389/fpls.2020.575829
  28. Gedeon, S., Ioannou, A., Balestrini, R., Fotopoulos, V., and Antoniou, C. 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
  29. 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.‏ https://doi.org/10.3390/agriculture11050463
  30. Haggag, I. A., Moustafa, M. M., Salama, A. N., Fadl, M. E., Drosos, M., Scopa, A., and Abd El-Raheem, A. A. 2024. Effect of Biostimulators as Foliar Application on Eggplant “Black Beauty Cultivar” Growth, Yield and Chemical Composition in Multi-Stressed Loamy Sand Soil. Horticulturae, 10(12), 1272. https://doi.org/10.3390/Horticulturae10121272
  31. Hariharan, G., Vathshalyan, N., Galahitigama, H., Wimalasiri, U., and Kumara, G. D. K. 2024. Potential of Foliar Application of Seaweed Extracts as a Biostimulant for Abiotic Stress Alleviation on Crop Production. Reviews in Agricultural Science, 12, 295-312.‏ https://doi.org/10.7831/ras.12.0_295
  32. Hancı, F., and Tuncer, G. 2020. How do foliar application of melatonin and L-tryptophan affect lettuce growth parameters under salt stress?. Turkish Journal of Agriculture-Food Science and Technology, 8(4), 960-964.‏ https://doi.org/10.24925/turjaf.v8i4.960-964.3224
  33. Ismail, M. A. M., Ahmad, M., Abdelrahman, T., Marwa, A., Sally, A., and Ali, M. 2024. Amino Acids and Polyamines in Foliar Spraying in Enhancing Medicinal Plant Resilience to Abiotic Stress–A Systematic Review. Integrative Biomedical Research (Former Journal of Angiotherapy), 8(11), 1-15.‏
  34. Kałużewicz, A., krzesiński, W., spiżewski, T., and Zaworska, A. 2017. Effect of biostimulants on several physiological characteristics and chlorophyll content in broccoli under drought stress and re-watering. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 45(1), 197-202.‏
  35. Kaya, G. 2025. A plant-derived biostimulant Aminolom Enzimatico® application stimulates chlorophyll content, electrolyte leakage, stomata density and root yield of radishes under salinity stress. PeerJ, 13, e18804.‏
  36. Kazemi, S., Pirmoradi, M. R., Raghami, M. and Malekzadeh, M. R. 2024. Enhancing the absorption of microelements by applying humic acid and zinc sulfate in Physalis alkekengi: Improve chlorophyll content and fruit quality. Greenhouse Plant Production Journal1(3), 68-82. https://doi.org/10.61186/gppj.1.3.68
  37. Khaleghi, G. 2024. The effect of hydropriming on germination characteristics and seedling growth of Nitraria schoberi seeds under salinity stress. Greenhouse Plant Production Journal1(1), 38-46. https://doi.org/10.61186/gppj.1.1.38
  38. Khan, M. A., Asaf, S., Khan, A. L., Jan, R., Kang, S. M., Kim, K. M., and Lee, I. J. 2020. Extending thermotolerance to tomato seedlings by inoculation with SA1 isolate of Bacillus cereus and comparison with exogenous humic acid application. PLoS One, 15(4), e0232228.‏ https://doi.org/10.1371/journal.pone.0232228
  39. Kiranmai, K., Lokanadha Rao, G., Pandurangaiah, M., Nareshkumar, A., Amaranatha Reddy, V., Lokesh, U., and Sudhakar, C. 2018. A novel WRKY transcription factor, MuWRKY3 (Macrotyloma uniflorum Lam. Verdc.) enhances drought stress tolerance in transgenic groundnut (Arachis hypogaea L.) plants. Frontiers in Plant Science, 9, 346.‏
  40. Kulkova, I., Wróbel, B., and Dobrzyński, J. 2024. Serratia spp. as plant growth-promoting bacteria alleviating salinity, drought, and nutrient imbalance stresses. Frontiers in Microbiology, 15, 1342331.‏ doi: 10.3389/fmicb.2024.1342331
  41. Lau, S. E., Lucas, W., Hamdan, M., Chan, C., Saidi, N., Ong-Abdullah, J., and Tan, B. 2025. Enhancing Plant Resilience to Abiotic Stress: The Power of Biostimulants. Phyton, 94(1), 1.‏ DOI:10.32604/phyton.2025.059930
  42. Liatile, P. C., Potgieter, G., and Moloi, M. J. 2022. A Natural Bio-Stimulant Consisting of a Mixture of Fish Protein Hydrolysates and Kelp Extract Enhances the Physiological, Biochemical and Growth Responses of Spinach under Different Water Levels. Plants, 11(23), 3374.‏
  43. Lin, Y., and Jones, M. L. 2022. Evaluating the growth-promoting effects of microbial biostimulants on greenhouse floriculture crops. HortScience, 57(1), 97-109.‏
  44. Ma, Y., Freitas, H., and Dias, M. C. 2022. Strategies and prospects for biostimulants to alleviate abiotic stress in plants. Frontiers in Plant Science, 13, 1024243.‏
  45. Mansori, L., Esna-Ashari, M., and Amerian, M. 2024. 'Effects of drought stress and mycorrhizal fungi on some characteristics of physiology and antioxidative enzymes in tarragon (Artemisia dracunculus L.), Plant Production and Genetics, 5(2), pp. 169-182. doi: 10.22034/plant.2024.141259.1103
  46. Moradtalab, N., Hajiboland, R., Aliasgharzad, N., Hartmann, T.E., and Neumann, G.2019. Silicon and the Association with an Arbuscular-Mycorrhizal Fungus (Rhizophagus clarus) Mitigate the Adverse Effects of Drought Stress on Str awberry. Agronomy, 9, 41. doi.org/10.3390/agronomy9010041
  47. Mordai, S., Amiri, J., Jabbarzadeh, Z., Rasouli-Sadaghiani, M., and Shaygan, A. 2024. Drench of humic acid mitigate the adverse impacts of alkalinity on rose. Ornamental Horticulture, 30, e242710.‏
  48. Muhie, S., Özdamar, C., Gökdaş, Z., Njie, E. S., Memiş, N., and Demir, İ. 2020. Effect of Solid Matrix Priming With Seaweed Extract on Germination and Seedling Performance of Onion Seeds under Abiotic Stress Conditions. Black Sea Journal of Agriculture, 3(4), 233-238.
  49. Muhie, S.H., Memis, N., Ozdamar, C., Gokdas, Z., Demir, I. 2021. Biostimulant priming for germination and seedling quality of carrot seeds under drought, salt and high temperature stress conditions. J. Agric. Environ. Food Sci., 5(3), 352-359 Doi: https://doi.org/10.31015/jaefs.2021.3.13
  50. Mukherjee, A., and Patel, J. S. 2020. Seaweed extract: biostimulator of plant defense and plant productivity. International Journal of Environmental Science and Technology, 17(1), 553-558. https://doi.org/10.1007/s13762-019-02442-z
  51. ‏Nezamdoost, D., Ghahremani, Z., Akbari, M.B., Barzegar, T., and Ranjbar, M. E. 2023. Can Seed Priming with Seaweed Extract Neutralize the Effects of Salinity on New Red Fire Leafy Lettuce Characteristics? . Gesunde Pflanzen 75, 955–969. https://doi.org/10.1007/s10343-022-00738-8
  52. Nordstedt, N. P., and Jones, M. L. 2020. Isolation of rhizosphere bacteria that improve quality and water stress tolerance in greenhouse ornamentals. Frontiers in Plant Science, 11, 826.‏
  53. Nordstedt, N. P., Chapin, L. J., Taylor, C. G., and Jones, M. L. 2020. Identification of Pseudomonas Spp. that increase ornamental crop quality during abiotic stress. Frontiers in plant science, 10, 1754.‏
  54. Ntanasi, T., Karavidas, I., Spyrou, G. P., Giannothanasis, E., Aliferis, K. A., Saitanis, C., and Ntatsi, G. 2024. Plant Biostimulants Enhance Tomato Resilience to Salinity Stress: Insights from Two Greek Landraces. Plants13(10), 1404.‏
  55. Pasbani, B., Salimi, A., Aliasgharzad, N., and Hajiboland, R. 2020. Colonization with arbuscular mycorrhizal fungi mitigates cold stress through improvement of antioxidant defense and accumulation of protecting molecules in eggplants. Scientia Horticulturae, 272, 109575.‏ https://doi.org/10.1016/j.scienta.2020.109575
  56. Pérez, L., Acosta, Y., Nápoles, L., Carvajal, C., Linares, C., Sershen, Lorenzo, J. C., Pérez, A. 2021. Pineapple stem-derived bromelain based priming improves pepper seed protein reserve mobilization, germination, emergence and plant growth. Physiology and Molecular Biology of Plants, 27(8), 1651-1657.‏ https://doi.org/10.1007/s12298-021-01038-7
  57. Piri, R., Moradi, A., Balouchi, H., and Salehi, A. 2019. Improvement of cumin (Cuminum cyminum) seed performance under drought stress by seed coating and biopriming. Scientia Horticulturae, 257, 108667.‏ https://doi.org/10.1016/j.scienta.2019.108667
  58. Petropoulos, S. A., Fernandes, Â., Plexida, S., Chrysargyris, A., Tzortzakis, N., Barreira, J. C. M., Barros, L., and Ferreira, I. C.F.R. 2020. Biostimulants application alleviates water stress effects on yield and chemical composition of greenhouse green bean (Phaseolus vulgaris L.). Agronomy, 10(2), 181.‏ https://doi.org/10.3390/agronomy10020181
  59. ‏Radwan, A. M., Ahmed, E. A., Donia, A. M., Mustafa, A. E., and Balah, M. A. 2023. Priming of Citrullus lanatus var. Colocynthoides seeds in seaweed extract improved seed germination, plant growth and performance under salinity conditions. Scientific Reports, 13(1), 11884.‏ https://doi.org/10.1038/s41598-023-38711-8
  60. Rahim, H. U., Ali, W., Uddin, M., Ahmad, S., Khan, K., Bibi, H., and Alatalo, J. M. 2024. Abiotic stresses in soils, their effects on plants, and mitigation strategies: a literature review. Chemistry and Ecology, 41(4), 552-585.‏ https://doi.org/10.1080/02757540.2024.2439830
  61. Raja, B., and Vidya, R. 2023. Application of seaweed extracts to mitigate biotic and abiotic stresses in plants. Physiology and Molecular Biology of Plants, 29(5), 641-661. https://doi.org/10.1007/s12298-023-01313-9
  62. Rajput, R. S., Singh, J., Singh, P., Vaishnav, A., and Singh, H. B. 2021. Influence of seed biopriming and vermiwash treatment on tomato plant's immunity and nutritional quality upon sclerotium rolfsii challenge inoculation. Journal of Plant Growth Regulation, 40, 1493-1509. https://doi.org/10.1007/s00344-020-10205-1
  63. Rezaei, Z., Roein, Z., Sabouri, A., and Hajinia, S. 2024a. Investigating the germination response of two Salvia species to water stress using the hydrotime model. Iranian Journal of Seed Sciences and Research, 11(2), 29-46. doi: 10.22124/jms.2024.8668
  64. Rezaei, Z., Roein, Z., Sabouri, A., and Hajinia, S. 2024b. Comparative analysis of seed germination in Thymus daenensis and T. vulgaris under water potential stress using hydrotime modelIranian Jurnal of seed Research. 11(1), 145-166. doi: 10.61186/yujs.11.1.145
  65. Roosta, H.R., Safarizadeh, M., Hamidpour, M., 2017. Effect of humic acid contained nano-fertile fertilizer spray on concentration of some nutrient elements in two lettuce cultivars in hydroponic system. J. Soil Plant Interact.-Isfahan Univ. Technol. 7, 51–59. 10.18869/acadpub.ejgcst.7.4.51
  66. Saddique, M., Kausar, A., IQRA, I., Akhter, N., Mujahid, N., Parveen, A., and Hussain, S. 2022. Amino acids application alleviated salinity stress in spinach (Spinacia oleracea L.) by improving oxidative defense, osmolyte
  67. Sales, J. R. D. S., Lacerda, C. F. D., Melo, A. S. D., Sousa, G. G. D., Mesquita, R. O., Ferreira, J. F. D. S., and Ribeiro, R. M. 2024. Algae biostimulant improves salt tolerance and visual quality of tropical ornamental herbaceous species. Revista Brasileira de Engenharia Agrícola e Ambiental, 28(4), e278645.‏
  68. Sangiorgio, D., Cellini, A., Donati, I., Pastore, C., Onofrietti, C., and Spinelli, F. 2020. Facing climate change: application of microbial biostimulants to mitigate stress in horticultural crops. Agronomy, 10(6), 794.‏
  69. Senousy, H. H., Hamoud, Y. A., Abu-Elsaoud, A. M., Mahmoud Al zoubi, O., Abdelbaky, N. F., Zia-Ur-Rehman, M., and Soliman, M. H. 2023. Algal bio-stimulants enhance salt tolerance in common bean: dissecting morphological, physiological, and genetic mechanisms for stress adaptation.  Plants, 12(21), 3714.‏
  70. Shahrajabian, M. H., Chaski, C., Polyzos, N., Tzortzakis, N., and Petropoulos, S. A. 2021. Sustainable agriculture systems in vegetable production using chitin and chitosan as plant biostimulants. Biomolecules, 11(6), 819.‏ https://doi.org/10.3390/biom11060819
  71. Shahrajabian, M. H., Cheng, Q., and Sun, W. 2022. The effects of amino acids, phenols and protein hydrolysates as biostimulants on sustainable crop production and alleviated stress. Recent Patents on Biotechnology, 16(4), 319-328.‏ DOI: 10.2174/1872208316666220412133749
  72. Sharafi, S. 2024. Increasing tolerance to salt-dryness stress of snail medic seedlings using Citationmagnetic field and ultrasonic waves. Greenhouse Plant Production Journal, 1(1): 27-37. https://doi.org/10.61186/gppj.1.1.27 
  73. Shirani Bidabadi, Siamak, and Mohammad Mehralian. 2020. "Arbuscular mycorrhizal fungi inoculation to enhance chilling stress tolerance of watermelon." Gesunde Pflanzen 72.2: 171-179.‏ https://doi.org/10.1007/s10343-020-00499-2
  74. Solgi, M., Bagnazari, M., Mohammadi, M., and Azizi, A. 2025. Thymbra spicata extract and arbuscular mycorrhizae improved the morphophysiological traits, biochemical properties, and essential oil content and composition of Rosemary (Rosmarinus officinalis L.) under salinity stress. BMC Plant Biology, 25(1), 220. https://doi.org/10.1186/s12870-025-06221-6
  75. Soroori, S., Danaee, E., Hemmati, K. and Ladan Moghadam, A. R. 2021. Effect of Foliar Application of Proline on Morphological and Physiological Traits of Calendula officinalis L. under drought Stress. J. Ornament. Plant., 11(1), 13-30. doi: 20.1001.1.22516433.2021.11.1.1.8
  76. Sorrentino, M., De Diego, N., Ugena, L., Spíchal, L., Lucini, L., Miras-Moreno, B., Zhang, L., Rouphael, Y., Colla, G., and Panzarová, K. 2021 Seed Priming With Protein Hydrolysates Improves Arabidopsis Growth and Stress Tolerance to Abiotic Stresses. Plant Science, 12, 626301. doi: 10.3389/fpls.2021.626301
  77. Soufi, H.R., Kalaji, H.M., Hamidpour, M and Malekzadeh, K. 2024. The roles of light in aCitation plant factory: photosynthesis efficiency and gas exchange parameters of lettuce as a function of light spectra. Greenhouse Plant Production Journal, 1(1): 1-26. https://doi.org/10.61186/gppj.1.1.1
  78. Sun, W., and Shahrajabian, M,H. 2023. The Application of Arbuscular Mycorrhizal Fungi as Microbial Biostimulant, Sustainable Approaches in Modern Agriculture. Plants.; 12(17):3101. https://doi.org/10.3390/Plants12173101
  79. Trovato, M., Funck, D., Forlani, G., Okumoto, S., and Amir, R. 2021. Amino acids in plants: Regulation and functions in development and stress defense. Frontiers in plant science, 12, 772810.‏
  80. ul Haq, S., Khan, A., Ali, M., Khattak, A. M., Gai, W. X., Zhang, H. X., and Gong, Z. H. 2019. Heat shock proteins: dynamic biomolecules to counter plant biotic and abiotic stresses. International Journal of Molecular Sciences, 20(21), 5321.‏ https://doi.org/10.3390/ijms20215321
  81. Van Oosten, M. J., Pepe, O., De Pascale, S., Silletti, S., and Maggio, A. 2017. The role of biostimulants and bioeffectors as alleviators of abiotic stress in crop plants. Chemical and Biological Technologies in Agriculture, 4, 1-12.
  82. Wazeer, H., Shridhar Gaonkar, S., Doria, E., Pagano, A., Balestrazzi, A., and Macovei, A. 2024. Plant-based biostimulants for seeds in the context of circular economy and sustainability. Plants, 13(7), 1004.‏ https://doi.org/10.3390/plants13071004
  83. Wróbel, M., Śliwakowski, W., Kowalczyk, P., Kramkowski, K., and Dobrzyński, J. 2023. Bioremediation of heavy metals by the genus Bacillus. International Journal of Environmental Research and Public Health, 20(6), 4964.‏ https://doi.org/10.3390/ijerph20064964
  84. Yadav, V. K., Jha, R. K., Kaushik, P., Altalayan, F. H., Al Balawi, T., and Alam, P. 2021. Traversing arbuscular mycorrhizal fungi and Pseudomonas fluorescens for carrot production under salinity. Saudi Journal of Biological Sciences, 28(8), 4217-4223.‏
  85. Yakob, B. K., Mam, E. L., and Sabirovich, G. M. 2024. Role of bio-stimulants on the advancement of vegetable production: A review. Agricultural Science and Technology (1313-8820), 16(4).‏
  86.  doi: 10.15547/ast.2024.04.035
  87. Yan, Z., Ma, T., Guo, S., Liu, R., and Li, M. 2021. Leaf anatomy, photosynthesis and chlorophyll fluorescence of lettuce as influenced by arbuscular mycorrhizal fungi under high temperature stress. Scientia Horticulturae, 280, 109933.‏
  88. Yusuf, R., Syakur, A., Mas’Ud, H., Latarang, B., Kartika, D. and Kristiansen, P. 2021, March. Application of local seaweed extracts to increase the growth and yield eggplant (Solanum melongena L.). In IOP Conference Series: Earth and Environmental Science (Vol. 681, No. 1, p. 012019). IOP Publishing. doi 10.1088/1755-1315/681/1/012019
  89. Zamljen, T., Medic, A., Veberic, R., Hudina, M., Grohar, M. C., and Slatnar, A. 2023. Influence of hydrolyzed animal protein-based biostimulant on primary, soluble and volatile secondary metabolism of Genovese and Greek-type basil grown under salt stress. Scientia Horticulturae, 319, 112178.‏
  90. Zhang, H., Li, Y., and Zhu, J. K. 2018. Developing naturally stress-resistant crops for a sustainable agriculture. Nat. Plants 4, 989–996. doi: 10.1038/s41477-018-0309-4
  91. Zulfiqar, F., Casadesús, A., Brockman, H., and 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
  92. Zuluaga, M. Y. A., Monterisi, S., Rouphael, Y., Colla, G., Lucini, L., Cesco, S., and Pii, Y. 2023. Different vegetal protein hydrolysates distinctively alleviate salinity stress in vegetable crops: A case study on tomato and lettuce. Frontiers in Plant Science, 14, 1077140.‏
  93. Zuzunaga-Rosas, J., Boscaiu, M., and Vicente, O. 2024. Agroindustrial By-Products as a source of biostimulants enhancing responses to abiotic stress of Horticultural crops. International Journal of Molecular Sciences, 25(6), 3525.‏ https://doi.org/10.3390/ijms25063525
Volume 2, Issue 1 - Serial Number 5
Winter 2025
Pages 121-136

  • Receive Date 20 February 2025
  • Revise Date 05 March 2025
  • Accept Date 20 March 2025
  • First Publish Date 30 March 2025
  • Publish Date 30 March 2025