The review summarizes and analyzes current scientific data on the toxic effects of heavy metals (HMs) and metalloids on the physiological, biochemical, molecular, and genetic processes in cereal crops (wheat, rice, maize, barley), and discusses innovative, environmentally friendly strategies to mitigate this stress. The mechanisms of HM phytotoxicity (Cd, Pb, As, Cr, Ni, Al), manifested in growth inhibition, destruction of the photosynthetic apparatus, disruption of ionic homeostasis, and generation of reactive oxygen species (ROS), are covered in detail. Central focus is given to the synergistic interaction of nanomaterials (inorganic, carbonaceous, and organic nanoparticles) with a complex phytohormonal ensemble, including abscisic, salicylic, and jasmonic acids, auxins, gibberellins, cytokinins, and brassinosteroids. The protective effects of presowing seed nano-priming and foliar application on plants are reviewed. The complex mechanisms of nanosystems action (particularly those based on ZnO, SiO2, TiO2, CeO2 Se, and chitosan) functioning as targeted hormone delivery agents, activating proton pumps, enhancing the barrier role of root cell walls, and stimulating the phenylpropanoid pathway for the biosynthesis of protective phenolic compounds and flavonoids are analyzed. At the molecular level, the role of the jasmonate signaling complex SCF-COI1-JAZ and the ability of combined nanoparticles and brassinosteroid application to significantly suppress the expression of HM transporter genes (OsHMA2, OsHMA6, OsNRAMP5) with simultaneous upregulation of the iron transporter OsIRT1 are highlighted. The prospects of using nanochelates and phytostimulants to minimize HM translocation into grain, reduce pesticide ecotoxicity, overcome nutrient deficiencies, and ensure global food security through sustainable agroecology systems are discussed.
Keywords: heavy metals, cereal crops, nanoparticles, food security, synergistic interaction, stress tolerance, phytohormones
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1. Rahman, S.U., Li, Y., Hussain, S., Hussain, B., Khan, W.-D., Riaz, L., Ashraf, M.N., Khaliq, M.A., Du, Z. & Cheng H. (2023). Role of phytohormones in heavy metal tolerance in plants: A review. Ecol. Indic., 146: 109844. https://doi.org/10.1016/j.ecolind.2022.109844
2. Ackova, D.G. (2018). Heavy metals and their general toxicity on plants. Plant Sci. Today, 5, No. 1, pp. 14-18. https://doi.org/10.14719/pst.2018.5.1.355
3. Vasilachi, I.C., Stoleru, V. & Gavrilescu, M. (2023). Analysis of heavy metal impacts on cereal crop growth and development in contaminated soils. Agriculture, 13, 1983. https://doi.org/10.3390/agriculture13101983
4. Paunov, M., Koleva, L., Vassilev, A., Vangronsveld, J. & Goltsev, V. (2018). Effects of different metals on photosynthesis: Cadmium and zinc affect chlorophyll fluorescence in durum wheat. Int. J. Mol. Sci., 19, No. 3, 787. https://doi.org/10.3390/ijms19030787
5. Riyazuddin, R., Nisha, N., Ejaz, B., Khan, M.I.R., Kumar, M., Ramteke, P.W. & Gupta, R. (2021). A comprehensive review on the heavy metal toxicity and sequestration in plants. Biomolecules, 12, No. 1, 43. https://doi.org/10.3390/biom12010043
6. Vodka, M.V. & Bilyavs'ka, N.O. (2016). Chloroplast structural and functional changes as biomarkers of heavy metal contamination. Biotachnol. Acta, 9, No 1, pp. 103-107. https://doi.org/10.15407/biotech9.01.103
7. Kosakivska, I.V., Babenko, L.M., Romanenko, K.O., Korotka, I.Y. & Potters, G. (2021). Molecular mechanisms of plant adaptive responses to heavy metals stress. Cell Biol. Int., 45, No. 2, pp. 258-272. https://doi.org/10.1002/cbin.11503
8. Bücker-Neto, L., Paiva, A.L.S., Machado, R.D., Arenhart, R.A. & Margis-Pinheiro, M. (2017). Interactions between plant hormones and heavy metals responses. Genet. Mol. Biol., 40, No. 1 suppl 1, pp. 373-386. https://doi.org/10.1590/1678-4685-gmb-2016-0087
9. Kosakivska, I.V., Babenko, L.M., Vasyuk, V.A. Voytenko, L.V. & Shcherbatiuk, M.M. (2024). Chapter "Natural Growth Regulators as Inducers of Resistance in Cereal Plants Against Extreme Environmental Factors" In book: "Regulation of adaptive responses in plants" Nova Science Publication, USA, p. 33-82. https://doi.org/10.52305/TXQB2084
10. Kosakivska, I., Babenko, L., Voytenko, L., Vasyuk, V., Shcherbatiuk, M. & Romanenko, K. (2025). Natural growth regulators in enhancing cereal crop resistance to lead contamination. Cereal Res. Commun., 53, No. 4, pp. 2063-2075. https://doi.org/10.1007/s42976-025-00697-6
11. Kosakivska, I.V., Vedenicheva, N.P., Babenko, L.M., Voytenko, L.V., Romanenko, K O. & Vasyuk, V.A. (2022). Exogenous phytohormones in the regulation of growth and development of cereals under abiotic stresses. Mol. Biol. Rep., 49, No. 1, pp. 617-628. https://doi.org/10.1007/s11033-021-06802-2
12. Vedenicheva, N. & Kosakivska, I. (2024). Chapter «The regulatory role of cytokinins in adaptive responses of cereal plants" In book: "Regulation of adaptive responses in plants" Nova Science Publication, USA, pp. 83-110. https://doi.org/10.52305/TXQB2084
13. Voytenko, L.V. & Kosakivska, I.V. (2025). Auxins as regulators of growth and development of cereal crops under abiotic stresses: a review. Adv. Biol. Earth Sci., 10, No. 1, pp. 5-31. https://doi.org/10.62476/abes.1015
14. Riyazuddin, R., Nisha, N. & Gupta, R. (2025). Phytonanotechnology in the mitigation of biotic and abiotic stresses in plants. Plant Nano Biol., 13, 100173. https://doi.org/10.1016/j.plana.2025.100173
15. Tripathi, D., Singh, M. & Rai, S.P. (2022). Crosstalk of nanoparticles and phytohormones regulate plant growth and metabolism under abiotic and biotic stress. Plant Stress, 6, 100107. https://doi.org/10.1016/j.stress.2022.100107
16. Ghorbani, A., Emamverdian, A., Pehlivan, N., Zargar, M., Razavi, S.M. & Chen, M. (2024). Nano-enabled agrochemicals: mitigating heavy metal toxicity and enhancing crop adaptability for sustainable crop production. J. Nanobiotechnology, 22, No. 1, 91. https://doi.org/10.1186/s12951-024-02371-1
17. Kekeli, M.A., Wang, Q. & Rui, Y. (2025). The role of nano-fertilizers in sustainable agriculture: Boosting crop yields and enhancing quality. Plants, 14, No. 4, 554. https://doi.org/10.3390/plants14040554
18. González-García, Y. & Juárez-Maldonado, A. (2025). Nanomaterials on plant growth and stress adaptation. Plants, 14, No. 11, 1651. https://doi.org/10.3390/plants14111651
19. Nazir, M.M., Li, G., Nawaz, M., Ahmed, T., Noman, M., Jalil, S. & Du, D. (2025). Nano-Enabled agrochemicals for heavy metal remediation in agriculture: Current status, mechanisms, and future prospects. Nanomaterials, 15, No. 20, 1588. https://doi.org/10.3390/nano15201588
20. Joudeh, N. & Linke, D. (2022). Nanoparticle classification, physicochemical properties, characterization, and applications: a comprehensive review for biologists. J. Nanobiotechnol., 20, 262 https://doi.org/10.1186/s12951-022-01477-8
21. Hussain, A., Rizwan, M., Ali, S., ur Rehman, M.Z., Qayyum, M.F., Nawaz, R. & Alyemeni, M.N. (2021). Combined use of different nanoparticles effectively decreased cadmium (Cd) concentration in grains of wheat grown in a field contaminated with Cd. Ecotoxicol. Environ. Saf., 215, 112139. https://doi.org/10.1016/j.ecoenv.2021.112139
22. Rehman, A., Khan, S., Sun, F., Peng, Z., Feng, K., Wang, N., Jia, Y., Pan, Z., He, S., Wang, L., Qayyum, A., Du, X. & Li, H. (2024). Exploring the nano-wonders: unveiling the role of Nanoparticles in enhancing salinity and drought tolerance in plants. Front. Plant Sci., 14, 1324176. https://doi.org/10.3389/fpls.2023.1324176
23. Wang, M., Luo, J., Li, H., Ge, C., Jing, F., Guo, J. & Zhou, D. (2025). Synergistic effect of foliar exposure to TiO2 nanoparticles and planting density modulates the metabolite profile and transcription to alleviate cadmium induced phytotoxicity to wheat (Triticum aestivum L.). Environ. Sci. Nano, 12, No. 1, pp. 879-893. https://doi.org/10.1039/D4EN00763H
24. Haghmadad Milani, M., Mohammadi, A., Panahirad, S., Farhadi, H., Labib, P., Kulak, M., Gohari, G., Fotopoulos, V. & Vita, F. (2024). Cerium oxide nanoparticles (CeO2 NPs) enhance salt tolerance in spearmint (Mentha spicata L.) by boosting the antioxidant system and increasing essential oil composition. Plants, 13, No. 20, 2934. https://doi.org/10.3390/plants13202934
25. Zhang, P., Jiang, Y., Schwab, F., Monikh, F.A., Grillo, R., White, J.C. & Lynch, I. (2024). Strategies for enhancing plant immunity and resilience using nanomaterials for sustainable agriculture. Environ. Sci. Technol., 58, No. 21, pp. 9051-9060. https://doi.org/10.1021/acs.est.4c03522
26. Oak, M.D. & Rajwade, J.M. (2026). Nanotechnology applications for enhancing stress tolerance in wheat: Past, present, and future prospects. In: Tiwari, R., Singh, B.K., Kumar, R. (eds) Wheat Stress Management. Springer, Singapore. https://doi.org/10.1007/978-981-92-0902-6_15
27. Morfín-Gutiérrez, A., García-Cerda, L.A., González-García, Y. & Juárez-Maldonado, A. (2025). Synthesis of Fe3O4@MCM-48 as nano fertilizer for growth stimulation in tomato plants. Plants, 14, No. 3, 405. https://doi.org/10.3390/plants14030405
28. Chen, F., Li, Y., Zia-ur-Rehman, M., Hussain, S.M., Qayyum, M.F., Rizwan, M., Alharby, H.F., Alabdallah, N.M., Alharbi, B.M. & Ali, S. (2023). Combined effects of zinc oxide nanoparticles and melatonin on wheat growth, chlorophyll contents, cadmium (Cd) and zinc uptake under Cd stress. Sci. Total Environ., 864, 161061. https://doi.org/10.1016/j.scitotenv.2022.161061
29. Lv, Z., Jiang, R., Chen, J. & Chen, W. (2020). Nanoparticle-mediated gene transformation strategies for plant genetic engineering. Plant Journal, 104, No. 4, pp. 880-891. https://doi.org/10.1111/tpj.14973
30. Hendler-Neumark, A. & Bisker, G. (2019). Fluorescent single-walled carbon nanotubes for protein detection. Sensors, 19, No. 24, 5403. https://doi.org/10.3390/s19245403
31. Kwak, S.Y., Lew, T.T.S., Sweeney, C.J., Koman, V.B., Wong, M.H., Bohmert-Tatarev, K., Snell, K.D., Seo, J.S., Chua, N.H. & Strano, M.S. (2019). Chloroplast-selective gene delivery and expression in planta using chitosan-complexed single-walled carbon nanotube carriers. Nat. Nanotechnol., 14, No. 5, pp. 447-455. https://doi.org/10.1038/s41565-019-0375-4
32. Demirer, G.S., Zhang, H., Goh, N.S., González-Grandío, E. & Landry, M.P. (2019). Carbon nanotube-mediated DNA delivery without transgene integration in intact plants. Nat. Protoc., 14, No. 10, pp. 2954-2971. https://doi.org/10.1038/s41596-019-0208-9
33. Demirer, G.S., Zhang, H., Goh, N.S., Pinals, R.L., Chang, R. & Landry, M.P. (2020). Carbon nanocarriers deliver siRNA to intact plant cells for efficient gene knockdown. Sci. Adv., 6, No. 26, eaaz0495. https://doi.org/10.1126/sciadv.aaz0495
34. Abd-Elsalam, K.A. (2024). Nanofertilizer delivery, effects and application methods. Elsevier, 486 p. https://doi.org/10.1016/C2022-0-02114-0
35. Adrees, M., Khan, Z.S., Hafeez, M., Rizwan, M., Hussain, K., Asrar, M., Alyemeni, M.N., Wijaya, L. & Ali, S. (2021). Foliar exposure of zinc oxide nanoparticles improved the growth of wheat (Triticum aestivum L.) and decreased cadmium concentration in grains under simultaneous Cd and water deficient stress. Ecotoxicol. Environ. Saf., 208, 111627. https://doi.org/10.1016/j.ecoenv.2020.111627
36. Zhao, L., Zhou, X., Kang, Z., Peralta-Videa, J.R. & Zhu, Y.G. (2024). Nano-enabled seed treatment: A new and sustainable approach to engineering climate-resilient crops. Sci. Total Environ., 910, 168640. https://doi.org/10.1016/j.scitotenv.2023.168640
37. Wu, T., Zhou, J. & Zhou, J. (2024). Comparison of soil addition, foliar spraying, seed soaking, and seed dressing of selenium and silicon nanoparticles effects on cadmium reduction in wheat (Triticum turgidum L.). Chemosphere, 362, 142681. https://doi.org/10.1016/j.chemosphere.2024.142681
38. Voytenko, L.V. & Kosakivska, I.V. (2016). Polyfunctional phytohormone abscisic acid. Bulletin of Kharkiv National Agrarian University. Series: Biology. 1, No. 37, pp. 27-41 [in Ukrainian]. https://repo.btu.kharkiv.ua/server/api/core/bitstreams/bec94a42-8d55-4b4b-9874-1e62c2a29421/content
39. Bano, A., Singh, K., Singh, S.P. & Sharma, P. (2023). Abscisic acid: Metabolism, signaling, and crosstalk with other phytohormones under heavy metal stress. Stresses, 3, pp. 665-686. https://doi.org/10.3390/stresses3040046
40. Kosakivska, I.V., Vasyuk, V.A., Voytenko, L.V. & Shcherbatiuk, M.M. (2022). Hormonal system of plants under the action of heavy metals. Kyiv.: M.G. Kholodny Institute of Botany, 176 p. [in Ukrainian]. https://www.botany.kiev.ua/doc/hormonal_monograph_2022.pdf
41. Huang, T.L., Nguyen, Q.T., Fu, S.F., Lin, C.Y., Chen, Y.C. & Huang, H.J. (2012). Transcriptomic changes and signalling pathway induced by arsenic stress in rice roots. Plant Mol. Biol., 80, pp. 587-608. https://doi.org/10.1007/s11103-012-9969-z
42. Lin, C.Y., Trinh, N.N., Lin, C.W. & Huang, H.J. (2013). Transcriptome analysis of phytohormone, transporters and signaling pathways in response to vanadium stress in rice roots. Plant Physiol Biochem. 66, 98-104. https://doi.org/10.1016/j.plaphy.2013.02.007
43. Khan, A.R., Salam, A., Li G., Iqbal, B., Ulhassan, Z., Liu, Q., Azhar, W., Liaquat, F., Hussain Shah, I., Shams ul Hassan, S. & Du, D. (2024). Nanoparticles and their crosstalk with stress mitigators: A novel approach towards abiotic stress tolerance in agricultural systems. Crop J., 12, No. 5, pp. 1280-1298. https://doi.org/10.1016/j.cj.2024.09.010
44. Zhao, Y., Wang, J., Huang, W., Zhang, D., Wu, J., Li, B., Li, M., Liu, L. & Yan, M. (2023). Abscisic-acid-regulated responses to alleviate cadmium toxicity in plants. Plants (Basel), 12, No. 5, 1023. https://doi.org/10.3390/plants12051023
45. Bunio, L.V., Tsvilynyuk, O.M. & Voytenko, L.V. (2025). The role of ABA in supporting the water balance in Carex hirta L. under conditions of oil polluted soil. Fìzìol. rosl. genet., 57, No. 2, pp. 167-180 [in Ukrainian]. https://doi.org/10.15407/frg2025.02.167
46. Kim, Y-H., Khan, A.L., Kim, D-H., Lee, S-Y., Kim, K-M., Waqas, M., Jung, H-Y., Shin, J-H., Kim, J-G. & Lee, I-J. (2014). Silicon mitigates heavy metal stress by regulating P-type heavy metal ATPases, Oryza sativa low silicon genes, and endogenous phytohormones. BMC Plant Biol., 14, pp. 13-20. https://doi.org/10.1186/1471-2229-14-13
47. Munzuroglu, Ö., Fikriye, K.Z. & Yahyagil, Z. (2008). The abscisic acid levels of wheat (Triticum aestivum L. cv. Çakmak 79) seeds that were germinated under heavy metal (Hg++, Cd++, Cu++) stress. G.U. J. Sci., 21, No. 1, pp. 1-7. https://scispace.com/pdf/the-abscisic-acid-levels-of-wheat-triticum-aestivum-l-cv-47qi9b9nbq.pdf
48. Zhao, X., Huang, S., Yao, Q., He, R., Wang, H., Xu, Z., Xing, W. & Liu, D. (2024). ABA-regulated MAPK signaling pathway promotes hormesis in sugar beet under cadmium exposure. J. Hazard. Mater., 480, 135968. https://doi.org/10.1016/j.jhazmat.2024.135968
49. Moya, J.L., Ros, R. & Picazo, I. (1995). Heavy metal-hormone interactions in rice plants: Effects on growth, net photosynthesis, and carbohydrate distribution. J. Plant Growth Regul., 14, pp. 61-67. https://doi.org/10.1007/BF00203115
50. Qin, M., Yan, J., Li, R., Jia, T., Sun, X., Liu, Z., El-Sheikh, M.A., Ahmad, P. & Liu, P. (2025). Integrated physiological, transcriptomic, and metabolomic investigation reveals that MgO NPs mediate the alleviation of cadmium stress in tobacco seedlings through ABA-regulated lignin synthesis. J. Hazard. Mater., 483, 136693. https://doi.org/10.1016/j.jhazmat.2024.136693
51. Ghouri, F., Sarwar, S., Sun, L., Riaz, M., Haider, F.U., Ashraf, H., Lai, M., Imran, M., Liu, J., Ali, S., Liu, X. & Shahid, M.Q. (2024). Silicon and iron nanoparticles protect rice against lead (Pb) stress by improving oxidative tolerance and minimizing Pb uptake. Sci. Rep., 14, No. 1, 5986. https://doi.org/10.1038/s41598-024-55810-2
52. İlhan, D. & Yazıcılar, B. (2025). Cd stress response in emmer wheat (Triticum dicoccum Schrank) varieties under in vitro conditions and remedial effect of CaO nanoparticles. Biology, 14, 394. https://doi.org/10.3390/biology14040394
53. Wahid, I., Kumari, S., Ahmad, R., Hussain, S.J., Alamri, S., Siddiqui, M.H. & Khan, M.I.R. (2020). Silver nanoparticle regulates salt tolerance in wheat through changes in ABA concentration, ion homeostasis, and defense systems. Biomolecules, 10, No. 11, 1506. https://doi.org/10.3390/biom10111506
54. Yue, L., Chen, F., Yu, K., Xiao, Z., Yu, X., Wang, Z. & Xing, B. (2019). Early development of apoplastic barriers and molecular mechanisms in juvenile maize roots in response to La2O3 nanoparticles. Sci. Total Environ., 653, pp. 675-683. https://doi.org/10.1016/j.scitotenv.2018.10.320
55. Sheteiwy, M.S., Fu, Y.F., Hu, Q.X., Nawaz, A., Guan, Y.J., Li, Z.H., Huang, Y.Y. & Hu, J.H. (2017). Regulation of ZnO nanoparticles-induced physiological and molecular changes by seed priming with humic acid in Oryza sativa seedlings. Plant Growth Regul., 83, No. 1, pp. 27-41. https://doi.org/10.1007/s10725-017-0281-4
56. Gui, X., Deng, Y., Rui, Y., Gao, B., Luo, W., Chen, S., Nhan, le V., Li, X., Liu, S., Han, Y., Liu, L. & Xing, B. (2015). Response difference of transgenic and conventional rice (Oryza sativa) to nanoparticles (gFe?O?). Environ. Sci. Pollut. Res., 22, No. 22, pp.17716-17723. https://doi.org/10.1007/s11356-015-4976-7
57. Yang, Z., Wei, H., Gan, Y., Liu, H., Cao, Y., An, H., Que, X., Gao, Y., Zhu, L., Tan, S., Liu, X. & Sun, L. (2025). Structural insights into auxin influx mediated by the Arabidopsis AUX1. Cell, 188, pp. 3960-3973. https://doi.org/10.1016/j.cell.2025.04.028
58. Janacek, D.P., Kolb, M., Schulz, L., Mergner, J., Kuster, B., Glanc, M., Friml, J., ten Tusscher, K., Schwechheimer, C. & Hammes, U.Z. (2024). Transport properties of canonical PIN-FORMED proteins from Arabidopsis and the role of the loop domain in auxin transport. Dev. Cell, 59, No. 24, pp. 3259-3271.e4. https://doi.org/10.1016/j.devcel.2024.09.020
59. Ung, K.L., Winkler, M., Schulz, L., Kolb, M., Janacek, D.P., Dedic, E., Stokes, D.L., Hammes, U.Z. & Pedersen, B.P. (2022). Structures and mechanism of the plant PIN-FORMED auxin transporter. Nature, 609, pp. 605-610. https://doi.org/10.1038/s41586-022-04883-y
60. Potters, G., Pasternak, T.P., Guisez, Y., Palme, K.J. & Jansen, M.A. (2007). Stress-induced morphogenic responses: growing out of trouble? Trends Plant Sci., 12, No. 3, pp. 98-105. https://doi.org/10.1016/j.tplants.2007.01.004
61. Pandya, P., Kumar, S., Patil, G., Mankad, M. & Shah, Z. (2024). Impact of ZnO nanopriming on physiological and biochemical traits of wheat (Triticum aestivum L.) seedling. CABI Agric Biosci, 5, 27. https://doi.org/10.1186/s43170-024-00228-z
62. Srivastav, A., Ganjewala, D., Singhal, R.K., Rajput, V.D., Minkina, T., Voloshina, M., Srivastava, S. & Shrivastava, M. (2021). Effect of ZnO nanoparticles on growth and biochemical responses of wheat and maize. Plants (Basel), 10, No. 12, 2556. https://doi.org/10.3390/plants10122556
63. Stałanowska, K., Szablińska-Piernik, J., Okorski, A. & Lahuta, L.B. (2023). Zinc oxide nanoparticles affect early seedlings' growth and polar metabolite profiles of pea (Pisum sativum L.) and wheat (Triticum aestivum L.). Int. J. Mol. Sci., 24, No. 19, 14992. https://doi.org/10.3390/ijms241914992
64. El-Badri, A.M., Batool, M., Wang, C., Hashem, A.M., Tabl, K.M., Nishawy, E., Kuai, J., Zhou, G. & Wang, B. (2021). Selenium and zinc oxide nanoparticles modulate the molecular and morpho-physiological processes during seed germination of Brassica napus under salt stress. Ecotoxicol. Environ. Saf., 225, 112695. https://doi.org/10.1016/j.ecoenv.2021.112695
65. Wei, J., Zou, Y., Li, P. & Yuan, X. (2020). Titanium dioxide nanoparticles promote root growth by interfering with auxin pathways in Arabidopsis thaliana. Phyton-Int. J. Exp. Bot., 89, No. 4, pp. 883-891. https://doi.org/10.32604/phyton.2020.010973
66. Sharma, A., Vishwakarma, K., Singh, N.K., Prakash, V., Ramawat, N., Prasad, R., Sahi, S., Singh, V.P., Tripathi, D.K. & Sharma, S. (2022). Synergistic action of silicon nanoparticles and indole acetic acid in alleviation of chromium (CrVI) toxicity in Oryza sativa seedlings. J. Biotechnol., 343, pp. 71-82. https://doi.org/10.1016/j.jbiotec.2021.09.005
67. Soleymani, S., Piri, S., Aazami, M.A. & Salehi, B. (2025). Cerium oxide nanoparticles alleviate drought stress in apple seedlings by regulating ion homeostasis, antioxidant defense, gene expression, and phytohormone balance. Sci. Rep., 15, 11805. https://doi.org/10.1038/s41598-025-96250-w
68. Jiang, M., Wang, J., Rui, M., Yang, L., Shen, J., Chu, H., Song, S. & Chen, Y. (2021). OsFTIP7 determines metallic oxide nanoparticles response and tolerance by regulating auxin biosynthesis in rice. J. Hazard. Mater., 403, 123946. https://doi.org/10.1016/j.jhazmat.2020.123946
69. Zhang, Z., Ke, M., Qu, Q., Peijnenburg, W.J.G.M., Lu, T., Zhang, Q., Ye, Y., Xu, P., Du, B., Sun, L. & Qian, H. (2018). Impact of copper nanoparticles and ionic copper exposure on wheat (Triticum aestivum L.) root morphology and antioxidant response. Environ. Pollut., 239, pp. 689-697. https://doi.org/10.1016/j.envpol.2018.04.066
70. Sponsel, V. & Hedden, P. (2010). Gibberellin Biosynthesis and Inactivation. Plant Hormones. Davies P.J. (Ed.). Springer, Dordrecht, 63-94. https://doi.org/10.1007/978-1-4020-2686-7_4
71. Vera-Sirera, F., Gomez, M.D. & Perez-Amador, M.A. (2016). Chapter 20 - DELLA Proteins, a group of GRAS transcription regulators that mediate gibberellin signaling. Editor(s): D.H. Gonzalez, Plant Transcription Factors. Academic Press, pp. 313-328. https://doi.org/10.1016/B978-0-12-800854-6.00020-8
72. Colebrook, E.H., Thomas, S.G., Phillips, A.L. & Hedden, P. (2014). The role of gibberellin signalling in plant responses to abiotic stress. J. Exp. Biol., 217. No. 1. pp. 67-75. https://doi.org/10.1242/jeb.089938
73. Kosakivska, I.V. & Vasyuk, V.A. (2021). Gibberelins in regulation of plant growth and development under abiotic stresses. Biotechnol. Acta, 14, No. 2. pp. 5-18. https://doi.org/10.15407/biotech14.02.005
74. Iftikhar, A., Ali, S., Yasmeen, T., Arif, M.S., Zubair, M., Rizwan, M., Alhaithloul, H.A.S., Alayafi, A.A.M. & Soliman, M.H. (2019). Effect of gibberellic acid on growth, photosynthesis and antioxidant defense system of wheat under zinc oxide nanoparticle stress. Environ. Pollut., 254, Pt B, 113109. https://doi.org/10.1016/j.envpol.2019.113109
75. Iftikhar, A., Rizwan, M., Adrees, M., Ali, S., Ur Rehman, M.Z., Qayyum, M.F. & Hussain, A. (2020). Effect of gibberellic acid on growth, biomass, and antioxidant defense system of wheat (Triticum aestivum L.) under cerium oxide nanoparticle stress. Environ. Sci. Pollut. Res., 27, No. 27, pp. 33809-33820. https://doi.org/10.1007/s11356-020-09661-9
76. Alharby, H.F., Rizwan, M., Iftikhar, A., Hussaini, K.M., Rehman, M.Z., Bamagoos, A.A., Alharbi, B.M., Asrar, M., Yasmeen, T. & Ali, S. (2021). Effect of gibberellic acid and titanium dioxide nanoparticles on growth, antioxidant defense system and mineral nutrient uptake in wheat. Ecotoxicol. Environ. Saf., 221, 112436. https://doi.org/10.1016/j.ecoenv.2021.112436
77. Al-Huqail, A.A., Alshehri, D., Nawaz, R., Irshad, M.A., Iftikhar, A., Hussaini, K.M., Rizwan, M., Alghanem, S.M.S. & Abeed, A.H.A. (2023). The effect of gibberellic acid on wheat growth and nutrient uptake under combined stress of cerium, zinc and titanium dioxide nanoparticles. Chemosphere, 336, 139199. https://doi.org/10.1016/j.chemosphere.2023.139199
78. Hakla, H.R., Sharma, S., Urfan, M., Yadav, N.S., Rajput, P., Kotwal, D., Abdel, Latef, A.A.H. & Pal, S. (2021). Gibberellins target shoot-root growth, morpho-physiological and molecular pathways to induce cadmium tolerance in Vigna radiata L. Agronomy, 11, No. 5, 896. https://doi.org/10.3390/agronomy11050896
79. da Cunha Neto, A.R., dos Santos Ambrósio, A., de Jesus Rodrigues Resende, A., Régis Santos, B. & Nadal, M.C. (2025). From cell division to stress tolerance: the versatile roles of cytokinins in plants. Phyton-Inter. J. Exp. Bot., 94, No. 3, pp. 539-560. https://doi.org/10.32604/phyton.2025.061776
80. Panozzo, A., Bolla, P.K., Barion, G., Botton, A. & Vamerali, T. (2025). Phytohormonal regulation of abiotic stress tolerance, leaf senescence and yield response in field crops: A comprehensive review. BioTech., 14, No. 1, 14. https://doi.org/10.3390/biotech14010014
81. Vankova, R., Landa, P., Podlipna, R., Dobrev, P.I., Prerostova, S., Langhansova, L., Gaudinova, A., Motkova, K., Knirsch, V. & Vanek, T. (2017). ZnO nanoparticle effects on hormonal pools in Arabidopsis thaliana. Sci. Total Environ., 593-594, pp. 535-542. https://doi.org/10.1016/j.scitotenv.2017.03.160
82. Tumburu, L., Andersen, C.P., Rygiewicz, P.T. & Reichman, J.R. (2015). Phenotypic and genomic responses to titanium dioxide and cerium oxide nanoparticles in Arabidopsis germinants. Environ. Toxicol. Chem., 34, No. 1, pp. 70-83. https://doi.org/10.1002/etc.2756
83. Vinković, T., Novák, O., Strnad, M., Goessler, W., Jurašin, D.D., Paradiković, N. & Vrček, I.V. (2017). Cytokinin response in pepper plants (Capsicum annuum L.) exposed to silver nanoparticles. Environ. Res., 156, pp. 10-18. https://doi.org/10.1016/j.envres.2017.03.015
84. Azhar, B.J., Noor, A., Zulfiqar, A., Zeenat, A., Ahmad, S. & Chishti, I. (2021). Effect of ZNO, SIO/sub 2/ and composite nanoparticles on arabidopsis thaliana and involvement of ethylene and cytokinin signaling pathways. Pak. J. Bot., 53, No. 2, pp. 437-446. https://inis.iaea.org/records/b4d7e-aj432 https://doi.org/10.30848/PJB2021-2(40)
85. Kosakivska, I.V. & Shcherbatiuk, M.M. (2025). Influence of lead pollution on the growth, development, and metabolism of cereal cops: the protective role of salicylic acid. Plant Physiol. Genet., 57, No. 1, pp. 3-17. https://doi.org/10.15407/frg2025.01.003
86. Liu, J., Qiu, G., Liu, C., Li, H., Chen, X., Fu, Q., Lin, Y. & Guo, B. (2022). Salicylic acid, a multifaceted hormone, combats abiotic stresses in plants. Life, 12, No. 6, 886. https://doi.org/10.3390/life12060886
87. Shcherbatiuk, M.M., Kosakivska, I.V., Voytenko, L.V. & Vasyuk V.A. (2026). Role of salicylic acid in activating cereal responses to heavy metal toxicity: mechanisms of action and quantitation methods. Fìzìol. rosl. genet. 58, No. 2, pp. 95-116 [in Ukrainian]. https://doi.org/10.15407/frg2026.02.095
88. Faizan, M., Sehar, S., Rajput, V.D., Faraz, A., Afzal, S., Minkina, T., Sushkova, S., Adil, M.F., Yu, F., Alatar, A.A., Akhter, F. & Faisal, M. (2021). Modulation of cellular redox status and antioxidant defense system after synergistic application of zinc oxide nanoparticles and salicylic acid in rice (Oryza sativa) plant under arsenic stress. Plants, 10, No. 11, 2254. https://doi.org/10.3390/plants10112254
89. Muhammad, S., Ulhassan, Z., Munir, R., Yasin, M.U., Islam, F., Zhang, K., Chen, W., Jan, M., Afzal, M., Muhammad, A., Hannan, F. & Zhou, W. (2025). Nanosilica and salicylic acid synergistically regulate cadmium toxicity in rice. Environ. Pollut., 364, Pt 1, 125331. https://doi.org/10.1016/j.envpol.2024.125331
90. Decsi, K., Ahmed, M., Abdul-Hamid, D., & Tóth, Z. (2025). The role of salicylic acid in activating plant stress responses - results of the past decade and future perspectives. Int. J. Mol. Sci., 26, No. 9, 4447. https://doi.org/10.3390/ijms26094447
91. Kour, J., Kohli, S.K., Khanna, K., Bakshi, P., Sharma, P., Singh, A.D., Ibrahim, M., Devi, K., Sharma, N., Ohri, P., Skalicky, M., Brestic, M., Bhardwaj, R., Landi, M. & Sharma, A. (2021). Brassinosteroid signaling, crosstalk and, physiological functions in plants under heavy metal stress. Front. Plant Sci., 12, 608061. https://doi.org/10.3389/fpls.2021.608061
92. Kothari, A. & Lachowiec, J. (2021). Roles of brassinosteroids in mitigating heat stress damage in cereal crops. Int. J. Mol. Sci., 22, No. 5, 2706. https://doi.org/10.3390/ijms22052706
93. Bartwal, A. & Arora, S. (2020). Brassinosteroids: Molecules with Myriad Roles. In: Mérillon, JM., Ramawat, K. (eds) Co-Evolution of Secondary Metabolites. Reference Series in Phytochemistry. Springer, Cham. https://doi.org/10.1007/978-3-319-96397-6_18
94. Nolan, T., Chen, J. & Yin, Y. (2017). Cross-talk of Brassinosteroid signaling in controlling growth and stress responses. Biochem. J., 474, pp. 2641-2661. https://doi.org/10.1042/BCJ20160633
95. Rattan, A., Kapoor, D., Kapoor, N., Bhardwaj, R. & Sharma, A. (2020). Brassinosteroids regulate functional components of antioxidative defense system in salt stressed maize seedlings. J. Plant Growth Regul., 4, pp. 1464-1475. https://doi.org/10.1007/s00344-020-10097-1
96. Wang, H., Feng, T., Peng, X., Yan, M., Zhou, P. & Tang, X. (2009). Ameliorative effects of brassinosteroid on excess manganese-induced oxidative stress in Zea mays L. leaves. Agr. Sci. China, 8, pp. 1063-1074. https://doi.org/10.1016/S1671-2927(08)60314-4
97. Bhardwaj, R., Arora, N., Sharma, P. & Arora, H.K. (2007). Effects of 28-homobrassinolide on seedling growth, lipid peroxzidation and antioxidative enzyme activities under nickel stress in seedlings of Zea mays L. Asian J. Plant Sc., 6, pp. 765-772. https://doi.org/10.3923/ajps.2007.765.772
98. Sharma, I., Kaur, N. & Pati, P.K. (2017). Brassinosteroids: A promising option in deciphering remedial strategies for abiotic stress tolerance in rice. Front. Plant Sci., 8, 2151. https://doi.org/10.3389/fpls.2017.02151
99. Zhou, Y., Xia, X., Yu, G., Wang, J., Wu, J., Wang, M., Yang, Y., Shi, K., Yu, Y., Chen, Z., Gan, J.& Yu, J. (2015). Brassinosteroids play a critical role in the regulation of pesticide metabolism in crop plants. Sci. Rep., 5, 9018. https://doi.org/10.1038/srep09018
100. Yusuf, M., Khan, T.A. & Fariduddin, Q. (2016). Interaction of epibrassinolide and selenium ameliorates the excess copper in Brassica juncea through altered proline metabolism and antioxidants. Ecotoxicol. Environ. Saf., 129, pp. 25-34. https://doi.org/10.1016/j.ecoenv.2016.03.001
101. Fan, L., Jiao, Q., Zhang, J., Zhang, H., Jiang, G-Y., Yang, J., Li, G., Fahad, S., Chen, Y., Agathokleous, E., Li, J., Jie, X., Liu, S. & Liu, H. (2026). Integrated morpho-physiological, biochemical and transcriptomic analyses reveal wheat responses to antimony toxicity and brassinosteroid-mediated alleviation. J. Hazard. Mater., 1, 00985. https://doi.org/10.1016/j.hazadv.2025.100985
102. Sehrish, A.K., Ahmad, S., Tabassam, R., Ai, F., Ali, S., Ali, S., Jamil, A., Naeem, M.K., Mahmood, Z. & Guo H. (2025). Zinc oxide nanoparticles and 24-epibrassinolide mitigate coexisting lead and cadmium toxicity in rice under elevated CO2: roles of metal transporters, antioxidant defense, and nutrient homeostasis Environ. Sci.: Nano, 12, 5322. https://doi.org/10.1039/D5EN00611B
103. Babenko, L.M., Kosakivska, I.V. & Skaterna, T.D. (2015). Jasmonic acid: Role in biotechnology and the regulation of plant biochemical processes. Biotechnol. Acta, 8, No. 2, pp. 36-51. https://doi.org/10.15407/biotech8.02.036
104. Sohn, S.I., Pandian, S., Rakkammal, K., Largia, M.J.V., Thamilarasan, S.K., Balaji, S., Zoclanclounon, Y.A.B., Shilpha, J. & Ramesh, M. (2022). Jasmonates in plant growth and development and elicitation of secondary metabolites: An updated overview. Front. Plant Sci., 13, 942789. https://doi.org/10.3389/fpls.2022.942789
105. Rehman, M., Saeed, M.S., Fan, X., Salam, A., Munir, R., Yasin, M.U., Khan, A.R., Muhammad, S., Ali, B., Ali, I., Khan, J. & Gan, Y. (2023). The Multifaceted Role of Jasmonic Acid in Plant Stress Mitigation: An Overview. Plants (Basel), 12, No. 23, 3982. https://doi.org/10.3390/plants12233982
106. Chen, X., Jiang, W., Tong, T., Chen, G., Zeng, F., Jang, S., Gao, W., Li, Z., Mak, M., Deng, F. & Chen, Z.H. (2021). Molecular interaction and evolution of jasmonate signaling with transport and detoxification of heavy metals and metalloids in plants. Front. Plant Sci., 12, 665842. https://doi.org/10.3389/fpls.2021.665842
107. Arya, S.S., Rookes, J.E., Cahill, D.M. & Lenka, S.K. (2022). Chitosan Nanoparticles and Their Combination with Methyl Jasmonate for the Elicitation of Phenolics and Flavonoids in Plant Cell Suspension Cultures. Int. J. Biol. Macromol., 214, pp. 632-641. https://doi.org/10.1016/j.ijbiomac.2022.06.145
108. Babenko, L.M., Smirnov, O.E., Romanenko, K.O., Trunova, O.K. & Kosakivska, I.V. (2019). Phenolic compounds in plants: biogenesis and functions. Ukr. Biochem. J., 91, No. 3, pp. 5-18. http://ukrbiochemjournal.org/wp-content/uploads/2019/05/Babenko_3_19.pdf https://doi.org/10.15407/ubj91.03.005
109. Martínez-Alesón García, P., García-Balboa, C., Romero-López, J., López-Rodas, V., Costas, E. & Baselga-Cervera, B. (2021). Fluctuation analysis to select for Samarium bio-uptaking microalgae clones the repurposing of a classical evolution experiment. Ecotox. Environ. Saf., 215, 112134. https://doi.org/10.1016/j.ecoenv.2021.112134
110. Zhao, J., Peng, J., Shang, C. & Yin, R. (2022). Revisiting the protocol for determining submicromolar concentrations of ozone in the water treated by advanced oxidation processes. Chemosphere, 303, Pt 2, 135117. https://doi.org/10.1016/j.chemosphere.2022.135117
111. Rodríguez-Seijo, A., Soares, C., Ribeiro, S., Amil, B.F., Patinha, C., Cachada, A., Fidalgo, F. & Pereira, R. (2022). Nano-Fe2O3 as a tool to restore plant growth in contaminated soils - Assessment of potentially toxic elements (bio)availability and redox homeostasis in Hordeum vulgare L. J. Hazar. Mater., 425, 127999. https://doi.org/10.1016/j.jhazmat.2021.127999
112. Yang, F. & Zhang, A. (2023). Involvement of METTL3 in arsenite-induced skin lesions by targeting the SOCS3/STAT3/Krt signaling pathway. Environ. Pollut., 316, Pt 2, 120634. https://doi.org/10.1016/j.envpol.2022.120634
113. Kim, Y-H., Khan, A.L., Waqas, M. & Lee, I-J. (2017). Silicon regulates antioxidant activities of crop plants under abiotic-induced oxidative stress: A review. Front. Plant Sci., 8, 510. https://doi.org/10.3389/fpls.2017.00510
114. Wei, T., Li, H., Wang, Y., Chi, M., Guo, J., Jia, H. & Zhang, C. (2025). Alleviation of cadmium toxicity and minimizing its accumulation in rice plants by methyl jasmonate: Performance and mechanisms. J. Biotechnol., 398, pp. 133-145. https://doi.org/10.1016/j.jbiotec.2024.12.009
115. Zhang, L., Bai, J., Zhai, Y., Zhang, K., Wang, Y., Xiao, R. & Jorquera, M.A. (2023). Effects of antibiotics on the endophyte and phyllosphere bacterial communities of lotus from above and below surface water in a typical shallow lake. Plant Physiol. Biochem., 201, 107812. https://doi.org/10.1016/j.plaphy.2023.107812
116. Jassim, A.Y., Karnwal, A., Pant, G., Gaur, A., Selvaraj, M., Dutta, J., Sachan, R.S.K., Kumar, M. & Nesterova, N. (2025). Harnessing nanotechnology for abiotic stress mitigation in agriculture: a sustainable approach to crop resilience. J. Plant Interact., 20, No. 1. https://doi.org/10.1080/17429145.2025.2586338
117. Li, Z., Zhong, Z., Zhang, B., Wang, W., Zhao, H., Seufitelli, G.V.S. & Resende, F.L.P. (2021). Microwave-assisted catalytic fast pyrolysis of rice husk over a hierarchical HZSM-5/MCM-41 catalyst prepared by organic base alkaline solutions. Sci. Total. Environ., 750, 141215. https://doi.org/10.1016/j.scitotenv.2020.141215
118. Leone, G., Pepi, S., Consumi, M., Lamponi, S., Fragai, M., Martinucci, M., Baldoneschi, V., Francesconi, O., Nativi, C. & Magnani, A. (2021). Sodium hyaluronate-g-2-((N-(6-aminohexyl)-4-methoxyphenyl)sulfonamido)-N-hydroxyacetamide with enhanced affinity towards MMP12 catalytic domain to be used as visco-supplement with increased degradation resistance. Carbohydr Polym., 271, 118452. https://doi.org/10.1016/j.carbpol.2021.118452
119. Kang, M., Liu, Y., Weng, Y., Wang, H. & Bai, X. (2024). A critical review on the toxicity regulation and ecological risks of zinc oxide nanoparticles to plants. Environ. Sci: Nano, 11, No. 1, pp. 14-35. https://doi.org/10.1039/D3EN00630A
120. Khan, S., Zahoor, M., Khan, R.S., Ikram, M. & Islam, N.U. (2023). The impact of silver nanoparticles on the growth of plants: The agriculture applications. Heliyon, 9, No. 6, e16928. https://doi.org/10.1016/j.heliyon.2023.e16928
121. Kumar, V., Dasgupta, N. & Ranjan, S. (Eds.). (2018). Environmental toxicity of nanomaterials. CRC Press. https://doi.org/10.1201/9781351252966
122. Singh, D. & Gurjar, B.R. (2022). Nanotechnology for agricultural applications: Facts, issues, knowledge gaps, and challenges in environmental risk assessment. J. Environ. Manag., 322, 116033. https://doi.org/10.1016/j.jenvman.2022.116033
123. Zaman, W., Ayaz, A. & Park, S. (2025). Nanomaterials in agriculture: A pathway to enhanced plant growth and abiotic stress resistance. Plants, 14, No. 5, 716. https://doi.org/10.3390/plants14050716
124. Lu, C., Hei R., Song, X., Fan, Z., Guo, D., Luo, J. & Ma, Y. (2023). Metal oxide nanoparticles inhibit nitrogen fixation and rhizosphere colonization by inducing ROS in associative nitrogen-fixing bacteria Pseudomonas stutzeri A1501. Chemosphere, 336, 139223. https://doi.org/10.1016/j.chemosphere.2023.139223
125. McGee, C.F. (2020). The effects of silver nanoparticles on the microbial nitrogen cycle: a review of the known risks. Environ. Sci. Pollut. Res., 27, No. 25, pp. 31061-31073. https://doi.org/10.1007/s11356-020-09548-9
126. Isibor, P.O., Oyegbade, S.A., Oni, J.G., Ahmed, W.W., Abiodun, E.O. & Oyewole, O.A. (2024). Nanoparticles in food chains: bioaccumulation and trophic transfer. In: Environmental Nanotoxicology: Combatting the Minute Contaminants (pp. 203-233). Cham: Springer Nature Switzerland. https://doi.org/10.1007/978-3-031-54154-4_11
127. Judy, J.D., Unrine, J.M. & Bertsch, P.M. (2011). Evidence for biomagnification of gold nanoparticles within a terrestrial food chain. Environ. Sci. Technol., 45, No. 2, pp. 776-781. https://doi.org/10.1021/es103031a
128. Li, M., Zhang, Y.Z., Feng, S., Zhang, X.X., Xi, Y.L. & Xiang, X.L. (2022). Bioaccumulation and biomagnification effects of nano-TiO2 in the aquatic food chain. Ecotoxicology, 31, pp. 1023-1034. https://doi.org/10.1007/s10646-022-02572-0
129. Kumari, R., Suman, K., Karmakar, S., Mishra, V., Lakra, S.G., Saurav, G.K. & Mahto, B.K. (2023). Regulation and safety measures for nanotechnology-based agri-products. Front. Genome Ed., 5, 1200987. https://doi.org/10.3389/fgeed.2023.1200987