Register      Login
Functional Plant Biology Functional Plant Biology Society
Plant function and evolutionary biology
RESEARCH ARTICLE (Open Access)

Green-synthesized ZnO and MgO nanoparticles modulate physiology and antioxidant defense in maize under alkaline stress

Muhammad Iftikhar https://orcid.org/0009-0008-6716-5367 A * and Anis Ali Shah https://orcid.org/0000-0003-2071-9939 A *
+ Author Affiliations
- Author Affiliations

A Department of Botany, Division of Science and Technology, University of Education, Lahore 54770, Pakistan.


Handling Editor: Honghong Wu

Functional Plant Biology 52, FP25200 https://doi.org/10.1071/FP25200
Submitted: 18 June 2025  Accepted: 4 September 2025  Published: 6 October 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)

Abstract

Alkaline stress severely impairs the growth and yield of Zea mays L. by disrupting physiological and biochemical functions. This study evaluated green-synthesized ZnO and MgO nanoparticles (NPs), prepared using neem and licorice extracts, for mitigating alkaline stress. NPs were nanosized, crystalline, and functionalized by phytochemicals, confirmed by scanning electron microscopy, FT-IR spectroscopy, UV-vis spectroscopy, and energy dispersive X-ray spectroscopy. A pot experiment using NPs (25–200 ppm) under control and alkaline stress assessed morphological, physiological, biochemical, and ionic responses. Alkaline stress reduced root fresh and dry weight to 2.60 and 0.66 g (−59.6%, −31.0%), shoot fresh and dry weight to 2.60 and 0.38 g (−59.6%, −70.0%), and chlorophyll a, b, and carotenoids to 1.31, 0.67, and 2.40 mg g−1 (−62.4%, −54.7%, −62.8%), whereas it increased malondialdehyde (MDA) (244.6%), H₂O₂ (457.7%), and relative membrane permeability (RMP) (55.9%). The combined ZnO (50 ppm) and MgO (50 ppm) treatment improved chlorophyll a, b, and carotenoids to 3.48, 1.48, and 6.45 mg g−1 (165.4%, 120.3%, 168.5%), and total soluble protein (392.8%), total protein (301.0%), proline (105.5%), glutathione (35.6%), and ascorbic acid (44.2%). Antioxidant enzymes increased, with superoxide dismutase at 29.52 U mg−1 (452.8%), peroxidase at 24.44 U mg−1 (862%), and ascorbate peroxidase at 51.62 U mg−1 (560%), whereas MDA, H2O2, and RMP (−78.1%) were reduced. High NP concentrations (ZnO 100 ppm + MgO 100 ppm) were toxic. Moderate ZnO and MgO NP doses enhanced resilience, yield stability, and sustainable agriculture.

Keywords: alkaline stress, antioxidant enzymes, MgONPs, plant physiology, stress markers, Zea mays L., ZnONPs.

References

Abbas Z, Hassan MA, Huang W, Yu H, Xu M, Chang X, Fang X, Liu L (2024) Influence of Magnesium Oxide (MgO) nanoparticles on maize (Zea mays L.). Agronomy 14(3), 617.
| Crossref | Google Scholar |

Abbasi Surki A, Nazari M, Fallah S, Iranipour R, Mousavi A (2020) The competitive effect of almond trees on light and nutrients absorption, crop growth rate, and the yield in almond–cereal agroforestry systems in semi-arid regions. Agroforestry Systems 94(3), 1111-1122.
| Crossref | Google Scholar |

Abdel Latef AA, Tran L-SP (2016) Impacts of priming with silicon on the growth and tolerance of maize plants to alkaline stress. Frontiers in Plant Science 7, 243.
| Crossref | Google Scholar |

Adil M, Bashir S, Bashir S, Aslam Z, Ahmad N, Younas T, Asghar RMA, Alkahtani J, Dwiningsih Y, Elshikh MS (2022) Zinc oxide nanoparticles improved chlorophyll contents, physical parameters, and wheat yield under salt stress. Frontiers in Plant Science 13, 932861.
| Crossref | Google Scholar |

Ahmad I, Ahmad B, Boote K, Hoogenboom G (2020) Adaptation strategies for maize production under climate change for semi-arid environments. European Journal of Agronomy 115, 126040.
| Crossref | Google Scholar |

Ahmad I, Ahmad W, Nepal J, Junaid MB, Bukhari NA, Usman M, Ahmad N, Khan RN (2024) Synergistic enhancement of maize crop yield and nutrient assimilation via zinc oxide nanoparticles and phosphorus fertilization. Journal of the Science of Food and Agriculture 104(11), 6733-6745.
| Crossref | Google Scholar | PubMed |

Ahmed N, Zhang B, Bozdar B, Chachar S, Rai M, Li J, Li Y, Hayat F, Chachar Z, Tu P (2023) The power of magnesium: unlocking the potential for increased yield, quality, and stress tolerance of horticultural crops. Frontiers in Plant Science 14, 1285512.
| Crossref | Google Scholar |

Ahmed AF, Fawzy M, Al-zahrani M, Abdelkader M (2025) Enhancing metabolic processes and water deficit stress tolerance of maize plants through biochar addition and foliar application of zinc nanoparticles. Russian Journal of Plant Physiology 72(1), 10.
| Crossref | Google Scholar |

Akhtar MS, Fiaz S, Aslam S, Chung S, Ditta A, Irshad MA, et al. (2024) Green synthesis of magnetite iron oxide nanoparticles using Azadirachta indica leaf extract loaded on reduced graphene oxide and degradation of methylene blue. Scientific Reports 14(1), 18172.
| Crossref | Google Scholar |

Akram Z, Hussain S, Mansoor M, Afzal M, Waqar A, Shabbir I (2014) Soil fertility and salinity status of Muzaffargarh District, Punjab Pakistan. Universal Journal of Agricultural Research 2, 242-249.
| Crossref | Google Scholar |

Al-Harbi LM, Ezzeldien M, Elhenawy AA, Said AH (2024) Assessment of the bioactivity of bioinspired magnesium oxide nanoparticles from the Azadirachta indica extract. Frontiers in Bioengineering and Biotechnology 12, 1480694.
| Crossref | Google Scholar |

Al-Kordy HMH, Sabry SA, Mabrouk MEM (2021) Statistical optimization of experimental parameters for extracellular synthesis of zinc oxide nanoparticles by a novel haloalaliphilic Alkalibacillus sp.W7. Scientific Reports 11, 10924.
| Crossref | Google Scholar |

Al-Selwey WA, Alsadon AA, Ibrahim AA, Labis JP, Seleiman MF (2023) Effects of zinc oxide and silicon dioxide nanoparticles on physiological, yield, and water use efficiency traits of potato grown under water deficit. Plants 12(1), 218.
| Crossref | Google Scholar |

Alabdallah NM, Alzahrani HS (2020) The potential mitigation effect of ZnO nanoparticles on [Abelmoschus esculentus L. Moench] metabolism under salt stress conditions. Saudi Journal of Biological Sciences 27(11), 3132-3137.
| Crossref | Google Scholar | PubMed |

Alexakis D, Gotsis D, Giakoumakis S (2012) Assessment of drainage water quality in pre- and post-irrigation seasons for supplemental irrigation use. Environmental Monitoring and Assessment 184, 5051-5063.
| Crossref | Google Scholar | PubMed |

Ali S, Ulhassan Z, Ali S, Kaleem Z, Yousaf MA, Sheteiwy MS, Ali S, Waseem M, Jalil S, Wang J, Zhou W (2024) Differential responses of Brassica napus cultivars to dual effects of magnesium oxide nanoparticles. Environmental Science and Pollution Research 31(8), 12446-12466.
| Crossref | Google Scholar | PubMed |

Almeida DM, Oliveira MM, Saibo NJM (2017) Regulation of Na+ and K+ homeostasis in plants: towards improved salt stress tolerance in crop plants. Genetics and Molecular Biology 40(1 suppl 1), 326-345.
| Crossref | Google Scholar | PubMed |

Alves TEP, Diniz AGA, Safadi GMVV, Silva-Neto CM (2024) Absorption of commercial and nanoparticulate ZnO and MgO synthesized by combustion reaction applied to maize soil. Environmental Nanotechnology, Monitoring & Management 22, 101005.
| Crossref | Google Scholar |

Amin ZS, Afzal M, Ahmed N, Zeshan B, Hashim NHHN, Yean CY (2023) Synthesis, characterization and biological activities of zinc oxide nanoparticles derived from secondary metabolites of Lentinula edodes. Molecules 28(8), 3532.
| Crossref | Google Scholar |

An Y, Gao Y, Tong S, Liu B (2021) Morphological and physiological traits related to the response and adaption of Bolboschoenus planiculmis seedlings grown under salt-alkaline stress conditions. Frontiers in Plant Science 12, 567782.
| Crossref | Google Scholar |

Arnon DI (1949) Copper enzymes in isolated chloroplasts. Polyphenoloxidase in Beta vulgaris. Plant Physiology 24, 1-15.
| Crossref | Google Scholar | PubMed |

Arrobas M, Afonso S, Ferreira IQ, Moutinho-Pereira J, Correia CM, Rodrigues MÂ (2017) Liming and application of nitrogen, phosphorus, potassium, and boron on a young plantation of chestnut. Turkish Journal of Agriculture and Forestry 41(6), 441-451.
| Crossref | Google Scholar |

Azarin K, Usatov A, Minkina T, Plotnikov A, Kasyanova A, Fedorenko A, Duplii N, Vechkanov E, Rajput VD, Mandzhieva S, Alamri S (2022) Effects of ZnO nanoparticles and its bulk form on growth, antioxidant defense system and expression of oxidative stress related genes in Hordeum vulgare L. Chemosphere 287, 132167.
| Crossref | Google Scholar |

Baloch Q, Kubar KA, Korai PK, Kalhoro SA, Junaid MB, Baloch MA, Zia-ul-hassan , Alkahtani J, Abrar M (2025) Evaluating vertical distribution of soil salinity patterns across multiple soil depths in a semi-arid dry region. Journal of Soil Science and Plant Nutrition 25, 5173-5185.
| Crossref | Google Scholar |

Barzegar M, Ahmadvand D, Sabouri Z, Darroudi M (2024) Phytoextract-mediated synthesis of magnesium oxide nanoparticles using Caccinia macranthera extract and examination of their photocatalytic and anticancer effects. Materials Research Bulletin 169, 112514.
| Crossref | Google Scholar |

Bates LS, Waldren RP, Teare ID (1973) Rapid determination of free proline for water-stress studies. Plant and Soil 39, 205-207.
| Crossref | Google Scholar |

Bayat M, Zargar M, Murtazova KM-S, Nakhaev MR, Shkurkin SI (2022) Ameliorating seed germination and seedling growth of nano-primed wheat and flax seeds using seven biogenic metal-based nanoparticles. Agronomy 12(4), 811.
| Crossref | Google Scholar |

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, 248-254.
| Crossref | Google Scholar | PubMed |

Broadley MR, White PJ (2010) Eats roots and leaves. Can edible horticultural crops address dietary calcium, magnesium and potassium deficiencies? Proceedings of the Nutrition Society 69(4), 601-612.
| Crossref | Google Scholar | PubMed |

Cakmak I, Horst WJ (1991) Effect of aluminium on lipid peroxidation, superoxide dismutase, catalase, and peroxidase activities in root tips of soybean (Glycine max). Physiologia Plantarum 83(3), 463-468.
| Crossref | Google Scholar |

Chance B, Maehly AC (1955) Assay of catalases and peroxidases. Methods in Enzymology 2, 764-775.
| Crossref | Google Scholar |

Chatepa LEC, Mwamatope B, Chikowe I, Masamba KG (2024) Effects of solvent extraction on the phytoconstituents and in vitro antioxidant activity properties of leaf extracts of the two selected medicinal plants from Malawi. BMC Complementary Medicine and Therapies 24(1), 317.
| Crossref | Google Scholar |

Dai F, Zhuang Q, Huang G, Deng H, Zhang X (2023) Infrared spectrum characteristics and quantification of OH groups in coal. ACS Omega 8(19), 17064-17076.
| Crossref | Google Scholar | PubMed |

Dangana RS, George RC, Shittu UO, Agboola FK (2023) Facile biosynthesis, characterisation and biotechnological application of ZnO nanoparticles mediated by leaves of Cnidoscolus aconitifolius. Artificial Cells, Nanomedicine, and Biotechnology 51(1), 309-317.
| Crossref | Google Scholar | PubMed |

de Sousa Ferreira L, de Souza Oliveira V, de Paula Marchiori JJ, Ferreira TC, Bernabé ACB, Boone GTF, dos Santos Pereira LL, Carriço E (2023) The nutrient magnesium in soil and plant: a review. International Journal of Plant & Soil Science 35(8), 136-144.
| Crossref | Google Scholar |

Deepak TS, Jayadeep PA (2022) Prospects of maize (corn) wet milling by-products as a source of functional food ingredients and nutraceuticals. Food Technology and Biotechnology 60(1), 109-120.
| Crossref | Google Scholar |

Dionisio-Sese ML, Tobita S (1998) Antioxidant responses of rice seedlings to salinity stress. Plant Science 135(1), 1-9.
| Crossref | Google Scholar |

Du J, AL-Huqail A, Cao Y, Yao H, Sun Y, Garaleh M, El Sayed Massoud E, Ali E, Assilzadeh H, Escorcia-Gutierrez J (2024) Green synthesis of zinc oxide nanoparticles from Sida acuta leaf extract for antibacterial and antioxidant applications, and catalytic degradation of dye through the use of convolutional neural network. Environmental Research 258, 119204.
| Crossref | Google Scholar |

Dutta S, Sinelshchikova A, Andreo J, Wuttke S (2024) Nanoscience and nanotechnology for water remediation: an earnest hope toward sustainability. Nanoscale Horizons 9(6), 885-899.
| Crossref | Google Scholar | PubMed |

Eissa D, Hegab RH, Abou-Shady A, Kotp YH (2022) Green synthesis of ZnO, MgO and SiO2 nanoparticles and its effect on irrigation water, soil properties, and Origanum majorana productivity. Scientific Reports 12(1), 5780.
| Crossref | Google Scholar |

El-Saber Batiha G, Magdy Beshbishy A, El-Mleeh A, M. Abdel-Daim M, Prasad Devkota H (2020) Traditional uses, bioactive chemical constituents, and pharmacological and toxicological activities of Glycyrrhiza glabra L.(Fabaceae). Biomolecules 10(3), 352.
| Crossref | Google Scholar |

Elbasiouny H, Elbehiry F, El-Ramady H (2022) Toxic effects of nanoparticles under combined stress on plants. In ‘Toxicity of nanoparticles in plants’. (Eds VD Rajput, T Minkina, S Sushkova, SS Mandzhieva, C Rensing) pp. 109–129. (Elsevier)

Faisal S, Naqvi S, Ali M, Lin L (2022) Comparative study of multifunctional properties of synthesised ZnO and MgO NPs for textiles applications. Pigment & Resin Technology 51(3), 301-308.
| Crossref | Google Scholar |

Faizan M, Faraz A, Yusuf M, Khan ST, Hayat S (2018) Zinc oxide nanoparticle-mediated changes in photosynthetic efficiency and antioxidant system of tomato plants. Photosynthetica 56, 678-686.
| Crossref | Google Scholar |

Faizan M, Alam P, Rajput VD, Faraz A, Afzal S, Ahmed SM, Yu F-Y, Minkina T, Hayat S (2023) Nanoparticle mediated plant tolerance to heavy metal stress: what we know? Sustainability 15(2), 1446.
| Crossref | Google Scholar |

Fernandes EA, dos Anjos Soares LA, de Lima GS, de Sousa Silva Neta AM, Roque IA, da Silva FA, et al. (2021) Cell damage, gas exchange, and growth of Annona squamosa L. under saline water irrigation and potassium fertilization. Semina: Ciências Agrárias, Londrina 42(3), 999-1018.
| Crossref | Google Scholar |

Gautam A, Sharma P, Ashokhan S, Yaacob JS, Kumar V, Guleria P (2023) Inhibitory impact of MgO nanoparticles on oxidative stress and other physiological attributes of spinach plant grown under field condition. Physiology and Molecular Biology of Plants 29(12), 1897-1913.
| Crossref | Google Scholar | PubMed |

Ghaffari Yaichi Z, Hassanpouraghdam MB, Rasouli F, Aazami MA, Vojodi Mehrabani L, Jabbari SF, Asadi M, Esfandiari E, Jimenez-Becker S (2025) Zinc oxide nanoparticles foliar use and arbuscular mycorrhiza inoculation retrieved salinity tolerance in Dracocephalum moldavica L. by modulating growth responses and essential oil constituents. Scientific Reports 15(1), 492.
| Crossref | Google Scholar |

Giannopolitis CN, Ries SK (1977) Superoxide dismutases: I. Occurrence in higher plants. Plant Physiology 59(2), 309-314.
| Crossref | Google Scholar | PubMed |

Girotti AW, Korytowski W (2023) Trafficking of oxidative stress-generated lipid hydroperoxides: pathophysiological implications. Free Radical Research 57(2), 130-139.
| Crossref | Google Scholar | PubMed |

Gowtham HG, Shilpa N, Singh SB, Aiyaz M, Abhilash MR, Nataraj K, Amruthesh KN, Ansari MA, Alomary MN, Murali M (2024) Toxicological effects of nanoparticles in plants: mechanisms involved at morphological, physiological, biochemical and molecular levels. Plant Physiology and Biochemistry 210, 108604.
| Crossref | Google Scholar |

Gupta A, Bharati R, Kubes J, Popelkova D, Praus L, Yang X, Severova L, Skalicky M, Brestic M (2024) Zinc oxide nanoparticles application alleviates salinity stress by modulating plant growth, biochemical attributes and nutrient homeostasis in Phaseolus vulgaris L. Frontiers in Plant Science 15, 1432258.
| Crossref | Google Scholar |

Hasan MK, Ara I, Mondal MSA, Kabir Y (2021) Phytochemistry, pharmacological activity, and potential health benefits of Glycyrrhiza glabra. Heliyon 7(6), e07240.
| Crossref | Google Scholar |

Hasanuzzaman M (2020) ‘Plant ecophysiology and adaptation under climate change: mechanisms and perspectives I: general consequences and plant responses.’ (Springer Nature)

Hassan ZA, Sarwar G, Aftab M, Manzoor MZ, Zafar A, Shehzad I, Riaz A, Hussain S, Sattar A, Niaz A (2022) Noxious impact of sodium bicarbonate (NaHCO3) on soil properties and ionic composition of rice plants. Journal of Agricultural Research 60(3), 193-202.
| Google Scholar |

Hu D, Li R, Dong S, Zhang J, Zhao B, Ren B, Ren H, Yao H, Wang Z, Liu P (2022) Maize (Zea mays L.) responses to salt stress in terms of root anatomy, respiration and antioxidative enzyme activity. BMC Plant Biology 22(1), 602.
| Crossref | Google Scholar |

Ikhuoria EU, Uwidia IE, Otabor GO, Ifijen IH (2024) Comparative analysis of magnesium oxide nanoparticles biosynthesized from rubber seed shell and rubber leaf extracts. Biomedical Materials & Devices 2(2), 1078-1088.
| Crossref | Google Scholar |

Inam A, Javad S, Naseer I, Alam P, Almutairi ZM, Faizan M, Zauq S, Shah AA (2024) Efficacy of chitosan loaded zinc oxide nanoparticles in alleviating the drastic effects of drought from corn crop. Plant Stress 14, 100617.
| Crossref | Google Scholar |

Irigoyen JJ, Einerich DW, Sánchez-Díaz M (1992) Water stress induced changes in concentrations of proline and total soluble sugars in nodulated alfalfa (Medicago sativa) plants. Physiologia Plantarum 84(1), 55-60.
| Crossref | Google Scholar |

Jamali K, Sahito F, Shah Z-u-H, Talpur NA, Rajpar I, Babar SK, Talpur KH, Afzal J, Amur A, Leghari Z (2023) Spatial variability mapping of selected soil properties of District Tando Allahyar, Sindh, Pakistan. Journal of Applied Research in Plant Sciences 4, 625-636.
| Crossref | Google Scholar |

Jamil M, Akhtar N, Iqbal MM, Khan MUH, Muslim N, Qazi MA (2021) Indexing of physico-chemical variables and fertility status of district Sahiwal soils, Punjab, Pakistan. Soil & Environment 40(1), 95-101.
| Crossref | Google Scholar |

Jampilek J, Kralova K (2022) Advances in biologically applicable graphene-based 2D nanomaterials. International Journal of Molecular Sciences 23(11), 6253.
| Crossref | Google Scholar |

Javed T, Shabbir R, Hussain S, Ahmed S, Saqib M, Rajendran K, Arslan D, Özçinar AB, El Sabagh AE (2025) Nanotechnology: a tool for mitigating abiotic stresses in crop plants. In ‘Nanotechnology for agriculture’. (Eds A Kumar, AE Sabagh) pp. 59–75. (Apple Academic Press)

Johnson-Beebout SE, Lauren JG, Duxbury JM (2009) Immobilization of zinc fertilizer in flooded soils monitored by adapted DTPA soil test. Communications in Soil Science and Plant Analysis 40(11–12), 1842-1861.
| Crossref | Google Scholar |

Jones MM, Turner NC (1978) Osmotic adjustment in leaves of sorghum in response to water deficits. Plant Physiology 61(1), 122-126.
| Crossref | Google Scholar | PubMed |

Kaczorowski R, Forenc T, Hunia J, Górny J, Janiszewski M, Jurek J, Komorowski M, Kapciak A, Pelczarska A (2024) Magnesium and zinc as vital micronutrients enhancing athletic performance and recovery–a review. Quality in Sport 33, 56021.
| Crossref | Google Scholar |

Kaleem M, Shah AA, Usman S, Xu W, Alsahli AA (2025) Magnesium oxide nanoparticles improved drought resilience in Coriander sativum L. through lifting antioxidant level, redox balancing, and improving photosynthetic efficiency. ACS Omega 10, 32813-32828.
| Crossref | Google Scholar | PubMed |

Kamath SD, Arunkumar D, Avinash NG, Samshuddin S (2015) Determination of total phenolic content and total antioxidant activity in locally consumed food stuffs in Moodbidri, Karnataka, India. Advances in Applied Science Research 6(6), 99-102.
| Google Scholar |

Kangathara N, Sabari V, Saravanan L, Elangovan S (2022) Synthesis, characterization, and comparison of pure zinc oxide and magnesium-doped zinc oxide nanoparticles and their application on ethanol sensing activities. Journal of Nanomaterials 2022(1), 1769278.
| Crossref | Google Scholar |

Kanwal S, Ilyas N, Shabir S, Saeed M, Gul R, Zahoor M, Batool N, Mazhar R (2018) Application of biochar in mitigation of negative effects of salinity stress in wheat (Triticum aestivum L.). Journal of Plant Nutrition 41(4), 526-538.
| Crossref | Google Scholar |

Kaur K, Jayarambabu N, Rao KV (2022) Comparative study of chemo-bio synthesized mgo nanoparticle on maize seed germination. IOP Conference Series: Materials Science and Engineering 1225, 012045.
| Crossref | Google Scholar |

Khan KS, Naveed M, Qadir MF, Yaseen M, Siddiqui MH (2023) Bio-organically acidified product-mediated improvements in phosphorus fertilizer utilization, uptake and yielding of Zea mays in calcareous soil. Plants 12(17), 3072.
| Crossref | Google Scholar |

Kiran S, Albargi HB, Afzal G, Aimun U, Anjum MN, Qadir MB, Khaliq Z, Jalalah M, Irfan M, Abdullah MM (2023) Azadirachta indica-assisted green synthesis of magnesium oxide nanoparticles for degradation of Reactive Red 195 dye: a sustainable environmental remedial approach. Applied Water Science 13(10), 193.
| Crossref | Google Scholar |

Kumar N, Nigam A, Pathak AK (2022) Assessment of genetic variability and path analysis in carrot (Daucus carota L.) genotypes. International Journal of Horticulture and Food Science 4, 1-9.
| Crossref | Google Scholar |

Kumar K, Jaiswal A, Koppolu UMK, Kumar KRR (2024) Alkaline stress disrupts growth, biochemistry, and ion homeostasis of chickpea (Cicer arietinum L.) roots. Frontiers in Agronomy 6, 1497054.
| Crossref | Google Scholar |

Lalarukh I, Zahra N, Al Huqail AA, Amjad SF, Al-Dhumri SA, Ghoneim AM, Alshahri AH, Almutari MM, Alhusayni FS, Al-Shammari WB, Poczai P, Mansoora N, Ayman M, Abbas MHH, Abdelhafez AA (2022) Exogenously applied ZnO nanoparticles induced salt tolerance in potentially high yielding modern wheat (Triticum aestivum L.) cultivars. Environmental Technology & Innovation 27, 102799.
| Crossref | Google Scholar |

Law MY, Charles SA, Halliwell B (1983) Glutathione and ascorbic acid in spinach (Spinacia oleracea) chloroplasts. The effect of hydrogen peroxide and of paraquat. Biochemical Journal 210(3), 899-903.
| Crossref | Google Scholar | PubMed |

Leach KA, Braun DM (2016) Soluble sugar and starch extraction and quantification from maize (Zea mays) leaves. Current Protocols in Plant Biology 1(1), 139-161.
| Crossref | Google Scholar | PubMed |

Lian J, Cheng L, Zhai X, Wu R, Liu W, Pan J, Shohag MJI, Xin X, He Z, Yang X (2022) Foliar spray of combined metal-oxide nanoparticles alters the accumulation, translocation and health risk of Cd in wheat (Triticum aestivum L.). Journal of Hazardous Materials 440, 129857.
| Crossref | Google Scholar |

Liang Y, Liu H, Fu Y, Li P, Li S, Gao Y (2023) Regulatory effects of silicon nanoparticles on the growth and photosynthesis of cotton seedlings under salt and low-temperature dual stress. BMC Plant Biology 23(1), 504.
| Crossref | Google Scholar |

Liu Z, Yuan D, Qin X, He P, Fu Y (2023) Effect of Mg-modified waste straw biochar on the chemical and biological properties of acidic soils. Molecules 28(13), 5225.
| Crossref | Google Scholar |

Ma Q, Wang H, Wu E, Yuan Y, Feng Y, Zhao L, Feng B (2023) Comprehensive physiological, transcriptomic, and metabolomic analysis of the response of Panicum miliaceum L. roots to alkaline stress. Land Degradation & Development 34(10), 2912-2930.
| Crossref | Google Scholar |

Mai Y, Ren Y, Deng S, Ashraf U, Tang X, Duan M, Mo Z (2024) Influence of ZnO nanoparticles on early growth stage of fragrant rice at low temperature (LT) stress. Journal of Soil Science and Plant Nutrition 24(1), 1301-1317.
| Crossref | Google Scholar |

Mansoor S, Ali Wani O, Lone JK, Manhas S, Kour N, Alam P, Ahmad A, Ahmad P (2022) Reactive oxygen species in plants: from source to sink. Antioxidants 11(2), 225.
| Crossref | Google Scholar |

Mansour MMF, Salama KHA (2020) Proline and abiotic stresses: responses and adaptation. In ‘Plant ecophysiology and adaptation under climate change: mechanisms and perspectives II: mechanisms of adaptation and stress amelioration’. (Ed. M Hasanuzzaman) pp. 357–397. (Springer)

Mardi A, Mohajjel Shoja H, Mohajel Kazemi E (2022) Comparative study of growth responses, photosynthetic pigment content, and gene expression pattern in tobacco plants treated with ZnO nano and ZnO bulk particles. Journal of Nanoparticle Research 24(10), 208.
| Crossref | Google Scholar |

Masood N, Hudson-Edwards KA, Zafar T, Farooqi A (2023) Natural carbon mineralization and its control on the geochemical evolution of coal-based aquifers in the Salt Range, Punjab, Pakistan. Environmental Geochemistry and Health 45(10), 7033-7050.
| Crossref | Google Scholar | PubMed |

Mazher M, Ishtiaq M, Hamid B, Haq SM, Mazhar A, Bashir F, Mazhar M, Mahmoud EA, Casini R, Alataway A, Dewidar AZ, Elansary HO (2023) Biosynthesis and characterization of calcium oxide nanoparticles from Citrullus colocynthis fruit extracts; their biocompatibility and bioactivities. Materials 16(7), 2768.
| Crossref | Google Scholar |

Mierek-Adamska A, Kulasek M, Dąbrowska GB, Blindauer CA (2025) Type 4 plant metallothioneins–players in zinc biofortification? Biological Reviews 100, 1229-1249.
| Crossref | Google Scholar | PubMed |

Miller GL (1959) Use of dinitrosalicylic acid reagent for determination of reducing sugar. Analytical Chemistry 31(3), 426-428.
| Crossref | Google Scholar |

Mirrani HM, Noreen Z, Usman S, Shah AA, Mahmoud EA, Elansary HO, Aslam M, Waqas A, Javed T (2024) Magnesium nanoparticles extirpate salt stress in carrots (Daucus carota L.) through metabolomics regulations. Plant Physiology and Biochemistry 207, 108383.
| Crossref | Google Scholar |

Moustakas M, Dobrikova A, Sperdouli I, Hanć A, Moustaka J, Adamakis I-DS, Apostolova E (2024) Photosystem II tolerance to excess zinc exposure and high light stress in Salvia sclarea L. Agronomy 14(3), 589.
| Crossref | Google Scholar |

Murtaza G, Qadir AA, Farooqi ZUR, Murtaza B (2023) Nutrient depletion in salt-affected soils and yield reduction. In ‘Soil constraints and productivity’. (Eds N Bolan, MB Kirkham) pp. 239–254. (CRC Press)

Mustafa G, Komatsu S (2022) Proteomics of plant-nanoparticle interaction mechanism. In ‘Plant and nanoparticles’. (Ed. JT Chen) pp. 67–84. (Springer)

Mustafa G, Chaudhari SK, Manzoor M, Batool S, Hatami M, Hasan M (2024) Zinc oxide nanoparticles mediated salinity stress mitigation in Pisum sativum: a physio-biochemical perspective. BMC Plant Biology 24(1), 835.
| Crossref | Google Scholar |

Nadeem M, Nazer Khan M, Abbas G, Fatima Z, Iqbal P, Ahmed M, Ali Raza M, Rehman A, Ul Haq E, Hayat A, Ali M, Ahmad S (2022) Application of CSM-CANEGRO model for climate change impact assessment and adaptation for sugarcane in semi-arid environment of Southern Punjab, Pakistan. International Journal of Plant Production 16(3), 443-466.
| Crossref | Google Scholar |

Nakano Y, Asada K (1981) Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant and Cell Physiology 22(5), 867-880.
| Crossref | Google Scholar |

Ni Y, Yang T, Ma Y, Zhang K, Soltis PS, Soltis DE, et al. (2021) Soil pH determines bacterial distribution and assembly processes in natural mountain forests of eastern China. Global Ecology and Biogeography 30(11), 2164-2177.
| Crossref | Google Scholar |

Ntshanka NM, Ejidike IP, Mthunzi FM, Moloto MJ, Mubiayi KP (2020) Investigation into the phytochemical profile, antioxidant and antibacterial potentials of Combretum molle and Acacia mearnsii leaf parts. Biomedical and Pharmacology Journal 13(4), 1683-1694.
| Crossref | Google Scholar |

Ogunyemi SO, Zhang M, Abdallah Y, Ahmed T, Qiu W, Ali MA, et al. (2020) The bio-synthesis of three metal oxide nanoparticles (ZnO, MnO2, and MgO) and their antibacterial activity against the bacterial leaf blight pathogen. Frontiers in Microbiology 11, 588326.
| Crossref | Google Scholar |

Olasan JO, Uzoma AC, Thomas AT, Ndidiamaka A, Mary DD (2024) Effect of magnesium oxide (MgO) nanoparticles on the performance of black-seeded beniseed (Sesamum indicum L.). Acta Chemica Malaysia (ACMY) 8(2), 107-113.
| Crossref | Google Scholar |

Opsahl S, Benner R (1999) Characterization of carbohydrates during early diagenesis of five vascular plant tissues. Organic Geochemistry 30(1), 83-94.
| Crossref | Google Scholar |

Pare B, Barde VS, Solanki VS, Agarwal N, Yadav VK, Alam MM, Gacem A, Alsufyani T, Khedher NB, Park J-W, Park S, Jeon B-H (2022) Green synthesis and characterization of LED-Irradiation-responsive nano ZnO catalyst and photocatalytic mineralization of malachite green dye. Water 14, 3221.
| Crossref | Google Scholar |

Parusnath M, Naidoo Y, Singh M, Rihan H, Dewir YH (2023) Phytochemical composition of Combretum molle (R. Br. ex G. Don.) Engl. & Diels leaf and stem extracts. Plants 12(8), 1702.
| Crossref | Google Scholar |

Pérez-Labrada F, Espinoza-Acosta JL, Bárcenas-Santana D, García-León E, López-Pérez MC (2024) Underlying mechanisms of action to improve plant growth and fruit quality in crops under alkaline stress. In ‘Abiotic Stress in crop plants-ecophysiological responses and molecular approaches’. (Eds M Hasanuzzaman, K Nahar) (IntechOpen)

Proshad R, Li J, Sun G, Zheng X, Yue H, Chen G, Zhang S, Li Z, Zhao Z (2024) Field application of hydroxyapatite and humic acid for remediation of metal-contaminated alkaline soil. Environmental Science and Pollution Research 31(9), 13155-13174.
| Crossref | Google Scholar | PubMed |

Qu R, Liu N, Wen Q, Guo J, Ge F (2024) Molecular mechanism of dissolvable metal nanoparticles-enhanced CO2 fixation by algae: metal-chlorophyll synthesis. Environmental Pollution 349, 123987.
| Crossref | Google Scholar |

Rajput VD, Harish , Singh RK, Verma KK, Sharma L, Quiroz-Figueroa FR, et al. (2021) Recent developments in enzymatic antioxidant defence mechanism in plants with special reference to abiotic stress. Biology 10(4), 267.
| Crossref | Google Scholar |

Rasouli F, Asadi M, Hassanpouraghdam MB, Aazami MA, Ebrahimzadeh A, Kakaei K, Dokoupil L, Mlcek J (2022) Foliar application of ZnO-NPs influences chlorophyll fluorescence and antioxidants pool in Capsicum annum L. under salinity. Horticulturae 8(10), 908.
| Crossref | Google Scholar |

Rathod A, Verma NS (2023) Impact of abiotic stress on agronomical crops. Frontiers of Agronomy 27,.
| Google Scholar |

Raza MAS, Muhammad F, Farooq M, Aslam MU, Akhter N, Toleikienė M, Binobead MA, Ali MA, Rizwan M, Iqbal R (2025) ZnO-nanoparticles and stage-based drought tolerance in wheat (Triticum aestivum L.): effect on morpho-physiology, nutrients uptake, grain yield and quality. Scientific Reports 15(1), 5309.
| Crossref | Google Scholar |

Rizwan M, Ali S, Ali B, Adrees M, Arshad M, Hussain A, et al. (2019) Zinc and iron oxide nanoparticles improved the plant growth and reduced the oxidative stress and cadmium concentration in wheat. Chemosphere 214, 269-277.
| Crossref | Google Scholar | PubMed |

Roque IA, Soares LADA, Lima GSD, Lopes IAP, Silva LDA, Fernandes PD (2022) Biomass, gas exchange and production of cherry tomato cultivated under saline water and nitrogen fertilization. Revista Caatinga 35(3), 686-696.
| Crossref | Google Scholar |

Sachdev S, Ansari SA, Ansari MI, Fujita M, Hasanuzzaman M (2021) Abiotic stress and reactive oxygen species: generation, signaling, and defense mechanisms. Antioxidants 10, 277.
| Crossref | Google Scholar |

Segatto C, Souza CA, Fiori MA, Lajús CR, Silva LL, Riella HG (2023) Seed treatment with magnesium nanoparticles alters phenology and increases grain yield and mineral content in maize. Australian Journal of Crop Science 17(2), 165-178.
| Crossref | Google Scholar |

Seghir BB, Hima M, Moulatti F, Sahraoui I, Ben Amor I, Zeghoud S, et al. (2023) Exploring the antibacterial potential of green-synthesized MgO and ZnO nanoparticles from two plant root extracts. Nanomaterials 13(17), 2425.
| Crossref | Google Scholar |

Seleiman MF, Ahmad A, Alhammad BA, Tola E (2023a) Exogenous application of zinc oxide nanoparticles improved antioxidants, photosynthetic, and yield traits in salt-stressed maize. Agronomy 13(10), 2645.
| Crossref | Google Scholar |

Seleiman MF, Ahmad A, Alshahrani TS (2023b) Integrative effects of zinc nanoparticle and PGRs to mitigate salt stress in maize. Agronomy 13(6), 1655.
| Crossref | Google Scholar |

Shanmugam J, Sharmili Sundararaj A, Shanmugasundaram R, Ravichandran B, Mani M, Mohammed Riyaz SU, Dhayalan M, Cid-Samamed A, Simal-Gandara J (2023) Green preparation of bract extract (Musa acuminate) doped magnesium oxide nanoparticles and their bioefficacy. Applied Organometallic Chemistry 37(5), e7063.
| Crossref | Google Scholar |

Shi Y, Jin X, Ackah M, Amoako FK, Li J, Tsigbey VE, Li H, Cui Z, Sun L, Zhao C, Zhao C (2024) Comparative physio-biochemical and transcriptome analyses reveal contrasting responses to magnesium imbalances in leaves of mulberry (Morus alba L.) plants. Antioxidants 13(5), 516.
| Crossref | Google Scholar |

Silva AA, Sousa AMF, Furtado CRG, Carvalho NMF (2022) Green magnesium oxide prepared by plant extracts: synthesis, properties and applications. Materials Today Sustainability 20, 100203.
| Crossref | Google Scholar |

Singh A, Rajput VD, Lalotra S, Agrawal S, Ghazaryan K, Singh J, Minkina T, Rajput P, Mandzhieva S, Alexiou A (2024a) Zinc oxide nanoparticles influence on plant tolerance to salinity stress: insights into physiological, biochemical, and molecular responses. Environmental Geochemistry and Health 46(5), 148.
| Crossref | Google Scholar |

Singh A, Sengar RS, Rajput VD, Al-Ghzawi AL, Shahi UP, Ghazaryan K, Minkina T, Tawaha ARMA, Al Zoubi OM, Habeeb T (2024b) Impact of salinity stress and zinc oxide nanoparticles on macro and micronutrient assimilation: unraveling the link between environmental factors and nutrient uptake. Journal of Ecological Engineering 25(2), 1-9.
| Crossref | Google Scholar |

Singh KM, Jha AB, Dubey RS, Sharma P (2025) Nanoparticle-mediated mitigation of salt stress-induced oxidative damage in plants: insights into signaling, gene expression, and antioxidant mechanisms. Environmental Science: Nano 12, 2983-3017.
| Crossref | Google Scholar |

Song J, Yu S, Yang R, Xiao J, Liu J (2023) Opportunities for the use of selenium nanoparticles in agriculture. NanoImpact 31, 100478.
| Crossref | Google Scholar |

Sriramachandrasekharan MV, Priya NG, Manivannan R, Prakash M (2022) Ameliorative role of Silicon on osmoprotectants, antioxidant enzymes and growth of maize grown under alkaline stress. Silicon 14, 6577-6585.
| Crossref | Google Scholar |

Srivastav A, Ganjewala D, Singhal RK, Rajput VD, Minkina T, Voloshina M, Srivastava S, Shrivastava M (2021) Effect of ZnO nanoparticles on growth and biochemical responses of wheat and maize. Plants 10(12), 2556.
| Crossref | Google Scholar |

Šulinskienė J, Bernotienė R, Baranauskienė D, Naginienė R, Stanevičienė I, Kašauskas A, Ivanov L (2019) Effect of zinc on the oxidative stress biomarkers in the brain of nickel-treated mice. Oxidative Medicine and Cellular Longevity 2019(1), 8549727.
| Crossref | Google Scholar |

Syed S, Islam A, Shabeer M, Nadhman A, Ahmad F, Irfan N, Mehwish S, Khan A (2024) Biomedical applications of green synthesized zinc oxide and magnesium-doped zinc oxide nanoparticles using aqueous extract of ziziphus oxyphylla leaves. IEEE Transactions on NanoBioscience 23(3), 418-427.
| Crossref | Google Scholar | PubMed |

Tabande L, Sepehri M, Yasrebi J, Zarei M, Ghasemi-Fasaei R, Khatabi B (2021) The root endophytic fungus Serendipita indica decreased the phytotoxicity of zinc oxide nanoparticles to alfalfa (Medicago sativa L.). 10.21203/rs.3.rs-219701/v1

Taj A, Bibi H, Akbar WA, Rahim HU, Iqbal M, Ullah S (2023) Effect of poultry manure and NPK compound fertilizer on soil physicochemical parameters, NPK availability, and uptake by spring maize (Zea mays L.) in alkaline-calcareous soil. Gesunde Pflanzen 75(2), 393-403.
| Crossref | Google Scholar |

Taj H, Noreen Z, Aslam M, Usman S, Shah AA, Rafique M, Raja V, El-Sheikh MA (2024) Effects of SNP, MgSO4, and MgO-NPs foliar application on Spinacia oleracea L. growth and physio-biochemical responses under cadmium stress. Scientific Reports 14(1), 26687.
| Crossref | Google Scholar |

Tanweer T, Rana NF, Saleem I, Shafique I, Alshahrani SM, Almukhlifi HA, Alotaibi AS, Alshareef SA, Menaa F (2022) Dental composites with magnesium doped zinc oxide nanoparticles prevent secondary caries in the alloxan-induced diabetic model. International Journal of Molecular Sciences 23(24), 15926.
| Crossref | Google Scholar |

Tryfon P, Sperdouli I, Adamakis I-DS, Mourdikoudis S, Moustakas M, Dendrinou-Samara C (2023) Impact of coated zinc oxide nanoparticles on photosystem II of tomato plants. Materials 16(17), 5846.
| Crossref | Google Scholar |

Upadhyay S, Mantha AK, Dhiman M (2020) Glycyrrhiza glabra (Licorice) root extract attenuates doxorubicin-induced cardiotoxicity via alleviating oxidative stress and stabilising the cardiac health in H9c2 cardiomyocytes. Journal of Ethnopharmacology 258, 112690.
| Crossref | Google Scholar |

Velikova V, Yordanov I, Edreva A (2000) Oxidative stress and some antioxidant systems in acid rain-treated bean plants: protective role of exogenous polyamines. Plant Science 151(1), 59-66.
| Crossref | Google Scholar |

Verma Y, Singh SK, Jatav HS, Rajput VD, Minkina T (2022) Interaction of zinc oxide nanoparticles with soil: insights into the chemical and biological properties. Environmental Geochemistry and Health 44, 221-234.
| Crossref | Google Scholar | PubMed |

Wang C, Wang H, Li Y, Li Q, Yan W, Zhang Y, Wu Z, Zhou Q (2021) Plant growth-promoting rhizobacteria isolation from rhizosphere of submerged macrophytes and their growth-promoting effect on Vallisneria natans under high sediment organic matter load. Microbial Biotechnology 14, 726-736.
| Crossref | Google Scholar | PubMed |

Wang W, Zhang F, Sun L, Yang L, Yang Y, Wang Y, Siddique KHM, Pang J (2022a) Alkaline salt inhibits seed germination and seedling growth of canola more than neutral salt. Frontiers in Plant Science 13, 814755.
| Crossref | Google Scholar |

Wang J, Li Q, Zhang M, Wang Y (2022b) The high pH value of alkaline salt destroys the root membrane permeability of Reaumuria trigyna and leads to its serious physiological decline. Journal of Plant Research 135(6), 785-798.
| Crossref | Google Scholar |

Ward RE, Carpenter CE (2010) Traditional methods for mineral analysis. In ‘Food analysis’. (Ed. SS Nielsen) pp. 201–215. (Springer)

Weber J, Starchenko V, Yuan K, Anovitz LM, Ievlev AV, Unocic RR, Borisevich AY, Boebinger MG, Stack AG (2023) Armoring of MgO by a passivation layer impedes direct air capture of CO2. Environmental Science & Technology 57(40), 14929-14937.
| Crossref | Google Scholar | PubMed |

Wittschier N, Faller G, Hensel A (2009) Aqueous extracts and polysaccharides from liquorice roots (Glycyrrhiza glabra L.) inhibit adhesion of Helicobacter pylori to human gastric mucosa. Journal of Ethnopharmacology 125(2), 218-223.
| Crossref | Google Scholar | PubMed |

Wu F, Fang Q, Yan S, Pan L, Tang X, Ye W (2020) Effects of zinc oxide nanoparticles on arsenic stress in rice (Oryza sativa L.): germination, early growth, and arsenic uptake. Environmental Science and Pollution Research 27, 26974-26981.
| Crossref | Google Scholar | PubMed |

Yang G, Rhodes D, Joly RJ (1996) Effects of high temperature on membrane stability and chlorophyll fluorescence in glycinebetaine-deficient and glycinebetaine-containing maize lines. Functional Plant Biology 23(4), 437-443.
| Crossref | Google Scholar |

Yang S, Xu Y, Tang Z, Jin S, Yang S (2024) The impact of alkaline stress on plant growth and its alkaline resistance mechanisms. International Journal of Molecular Sciences 25(24), 13719.
| Crossref | Google Scholar |

Zhai J, Xian X, Zhang Z, Wang Y (2025) Nano-zinc oxide can enhance the tolerance of apple rootstock M9-T337 seedlings to saline alkali stress by initiating a variety of physiological and biochemical pathways. Plants 14(2), 233.
| Crossref | Google Scholar |

Zhang D, Tang Y, Zhang C, Huhe FNU, Wu B, Gong X, Chuang SSC, Zheng J (2022) Formulating zwitterionic, responsive polymers for designing smart soils. Small 18(38), 2203899.
| Crossref | Google Scholar |

Łukasik I, Sytykiewicz H, Goławska S (2021) Effect of the cereal aphid infestation on the oxidative damages of protein in the maize (Zea mays L.). Acta Scientiarum Polonorum Hortorum Cultus 20(1), 81-89.
| Crossref | Google Scholar |