{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T21:42:58Z","timestamp":1787953378732,"version":"build-2784847793"},"reference-count":81,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2025,6,9]],"date-time":"2025-06-09T00:00:00Z","timestamp":1749427200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Corps Science and Technology Development Special","award":["2018AB012"],"award-info":[{"award-number":["2018AB012"]}]},{"name":"Corps Science and Technology Development Special","award":["20240428"],"award-info":[{"award-number":["20240428"]}]},{"name":"Tianshan Talents Teaching Masters Project in Xinjiang","award":["2018AB012"],"award-info":[{"award-number":["2018AB012"]}]},{"name":"Tianshan Talents Teaching Masters Project in Xinjiang","award":["20240428"],"award-info":[{"award-number":["20240428"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Glycyrrhiza uralensis Fisch. is an important economic plant. With its wild populations on the brink of extinction and the area of salinized soil increasing sharply, farmers have gradually used saline soil to carry out artificial cultivation of the licorice. However, the salt stress has led to a significant decrease in the yield and quality of its medicinal organ (root), seriously restricting the sustainable development of the licorice industry. Therefore, we investigated zinc oxide nanoparticles (ZnO NPs) as a nano-fertilizer to enhance root biomass and bioactive compound accumulation under salinity. Our results indicate that under 160 mM NaCl stress, the application of 30 mg\/kg ZnO NPs increased the root biomass of the licorice and the contents of glycyrrhizic acid, glycyrrhizin, and total flavonoids in the roots by 182%, 158%, 87%, and 201%, respectively. And the ZnO treatment made the enzyme activities of SOD, CAT, and POD exhibit increase, and made the levels of superoxide anions, electrolyte leakage, soluble sugar, and proline reduce. These results demonstrate that ZnO NPs not only enhance salt tolerance but also redirect metabolic resources toward medicinal compound biosynthesis. Our findings provide a mechanistic basis for utilizing nanotechnology to sustainably cultivate the licorice in marginal saline environments, bridging agricultural productivity and pharmacological value.<\/jats:p>","DOI":"10.3390\/plants14121763","type":"journal-article","created":{"date-parts":[[2025,6,10]],"date-time":"2025-06-10T04:42:25Z","timestamp":1749530545000},"page":"1763","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["ZnO NPs: A Nanomaterial-Based Fertilizer That Significantly Enhanced Salt Tolerance of Glycyrrhiza uralensis Fisch and Improved the Yield and Quality of Its Root"],"prefix":"10.3390","volume":"14","author":[{"given":"Ning","family":"Wu","sequence":"first","affiliation":[{"name":"Ministry of Education Key Laboratory of Xinjiang Phytomedicine Resource Utilization, The Key Laboratory of Oasis Town and Mountain-Basin System Ecology, College of Life Sciences, Shihezi University, Shihezi 832003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7857-6761","authenticated-orcid":false,"given":"Miao","family":"Ma","sequence":"additional","affiliation":[{"name":"Ministry of Education Key Laboratory of Xinjiang Phytomedicine Resource Utilization, The Key Laboratory of Oasis Town and Mountain-Basin System Ecology, College of Life Sciences, Shihezi University, Shihezi 832003, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,6,9]]},"reference":[{"key":"ref_1","unstructured":"FAO (2024, December 15). Land & Water. Available online: https:\/\/www.fao.org\/land-water\/solaw2021\/facts\/zh\/."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Yue, J.Y., Wang, Y.J., Jiao, J.L., and Wang, H.Z. (2021). Silencing of ATG2 and ATG7 promotes programmed cell death in wheat via inhibition of autophagy under salt stress. Ecotoxicol. Environ. Saf., 225.","DOI":"10.1016\/j.ecoenv.2021.112761"},{"key":"ref_3","unstructured":"Caon, L., Forlano, N., Keene, C., Sala, M., Sorokin, A., Verbeke, I., and Ward, C. (2015). Global status of soil salinization and sodification. Status of the World\u2019s Soil Resources\u2014Main Report, Food and Agriculture Organization of the United Nations and Intergovernmental Technical Panel on Soils."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"He, P., Li, J., Yu, S., Ma, T., Ding, J., Zhang, F., Chen, K., Guo, S., and Peng, S. (2023). Soil moisture regulation under mulched drip irrigation influences the soil salt distribution and growth of cotton in southern Xinjiang, China. Plants, 12.","DOI":"10.3390\/plants12040791"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Yan, B., Hou, J., Li, W., Luo, L., Ye, M., Zhao, Z., and Wang, W. (2023). A review on the plant resources of important medicinal licorice. J. Ethnopharmacol., 301.","DOI":"10.1016\/j.jep.2022.115823"},{"key":"ref_6","first-page":"726","article-title":"Chemical profile and anti-inflammatory activity of total flavonoids from Glycyrrhiza uralensis Fisch","volume":"17","author":"Yin","year":"2018","journal-title":"Iran. J. Pharm. Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"826","DOI":"10.2174\/1389557519666190119111125","article-title":"Research progress of glycyrrhizic acid on antiviral activity","volume":"19","author":"Sun","year":"2019","journal-title":"Mini-Rev. Med. Chem."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Zhao, C., Wang, D., Gao, Z., Kan, H., Qiu, F., Chen, L., and Li, H. (2020). Licocoumarone induces BxPC-3 pancreatic adenocarcinoma cell death by inhibiting DYRK1A. Chem.-Biol. Interact., 316.","DOI":"10.1016\/j.cbi.2019.108913"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.taap.2017.04.015","article-title":"Licoricidin inhibits the growth of SW480 human colorectal adenocarcinoma cells in vitro and in vivo by inducing cycle arrest, apoptosis and autophagy","volume":"326","author":"Ji","year":"2017","journal-title":"Toxicol. Appl. Pharmacol."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Lu, J., Chang, T., Tang, X., Wang, Q., Pan, D., Wang, J., Nan, H., Zhang, W., and Liu, L. (2024). A bibliometric review of Glycyrrhizae Radix et Rhizoma (licorice) research: Insights and future directions. J. Ethnopharmacol., 321.","DOI":"10.1016\/j.jep.2023.117409"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Dong, X., Ma, X., Zhao, Z., and Ma, M. (2024). Exogenous betaine enhances salt tolerance of Glycyrrhiza uralensis through multiple pathways. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-024-04851-w"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Schmid, C., Brockhoff, A., Shoshan-Galeczki, Y.B., Kranz, M., Stark, T.D., Erkaya, R., Meyerhof, W., Niv, M.Y., Dawid, C., and Hofmann, T. (2021). Comprehensive structure-activity-relationship studies of sensory active compounds in licorice (Glycyrrhiza glabra). Food Chem., 364.","DOI":"10.1016\/j.foodchem.2021.130420"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"554","DOI":"10.1271\/bbb.58.554","article-title":"Sweetness of glycyrrhetic acid 3-O-beta-D-monoglucuronide and the related glycosides","volume":"58","author":"Mizutani","year":"1994","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Alagawany, M., Elnesr, S.S., Farag, M.R., Abd El-Hack, M.E., Khafaga, A.F., Taha, A.E., Tiwari, R., Yatoo, M.I., Bhatt, P., and Marappan, G. (2019). Use of licorice (Glycyrrhiza glabra) herb as a feed additive in poultry: Current knowledge and prospects. Animals, 9.","DOI":"10.3390\/ani9080536"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Liu, Y., Li, D., Gao, H., Li, Y., Chen, W., Jiao, S., and Wei, G. (2022). Regulation of soil micro-foodwebs to root secondary metabolites in cultivated and wild licorice plants. Sci. Total Environ., 828.","DOI":"10.1016\/j.scitotenv.2022.154302"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Yao, H., Wang, F., Bi, Q., Liu, H., Liu, L., Xiao, G., Zhu, J., Shen, H., and Li, H. (2022). Combined analysis of pharmaceutical active ingredients and transcriptomes of Glycyrrhiza uralensis under PEG6000-induced drought stress revealed glycyrrhizic acid and flavonoids accumulation via JA-mediated signaling. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.920172"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2192","DOI":"10.1007\/s42729-023-01542-7","article-title":"Unveiling mechanisms of silicon-induced salt or\/and drought tolerance in Glycyrrhiza uralensis Fisch. by physiological and transcriptomic analysis","volume":"24","author":"Fan","year":"2024","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1007\/s10725-023-01054-5","article-title":"Silicon improves the growth of Glycyrrhiza uralensis Fisch. under salt and\/or drought stress by regulating respiration metabolism","volume":"101","author":"Fan","year":"2023","journal-title":"Plant Growth Regul."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"25916","DOI":"10.1007\/s11356-018-2595-9","article-title":"Silicon improves salt tolerance of Glycyrrhiza uralensis Fisch. by ameliorating osmotic and oxidative stresses and improving phytohormonal balance","volume":"25","author":"Zhang","year":"2018","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Jia, T., Gu, J., Ma, M., and Song, Y. (2024). Lanthanum significantly contributes to the growth of the fine roots\u2019 morphology and phosphorus uptake efficiency by increasing the yield and quality of Glycyrrhiza uralensis taproots. Plants, 13.","DOI":"10.3390\/plants13040474"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Nishidono, Y., and Tanaka, K. (2025). Exogenous ethanol treatment promotes glycyrrhizin accumulation in aseptically grown Glycyrrhiza uralensis seedlings. Plant Signal. Behav., 20.","DOI":"10.1080\/15592324.2025.2472012"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1007\/s40011-016-0791-2","article-title":"Nanopesticides: Opportunities in crop protection and associated environmental risks","volume":"88","author":"Pandey","year":"2018","journal-title":"Proc. Natl. Acad. Sci. India Sect. B"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1515\/biol-2022-0461","article-title":"Agronomic and economic performance of mung bean (Vigna radiata L.) varieties in response to rates of blended NPS fertilizer in Kindo Koysha district, Southern Ethiopia","volume":"17","author":"Baza","year":"2022","journal-title":"Open Life Sci."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Rui, M., Ma, C., Hao, Y., Guo, J., Rui, Y., Tang, X., Zhao, Q., Fan, X., Zhang, Z., and Hou, T. (2016). Iron oxide nanoparticles as a potential iron fertilizer for peanut (Arachis hypogaea). Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.00815"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1007\/s11104-013-1967-7","article-title":"Uptake of zinc and phosphorus by plants is affected by zinc fertiliser material and arbuscular mycorrhizas","volume":"376","author":"Turney","year":"2014","journal-title":"Plant Soil"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1016\/j.jphotobiol.2016.12.011","article-title":"A review on biogenic synthesis of ZnO nanoparticles using plant extracts and microbes: A prospect towards green chemistry","volume":"166","author":"Ahmed","year":"2017","journal-title":"J. Photochem. Photobiol. B"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"362","DOI":"10.2174\/1389203721666201016144848","article-title":"Zinc oxide nanoparticles help to enhance plant growth and alleviate abiotic stress: A review","volume":"22","author":"Faizan","year":"2021","journal-title":"Curr. Protein Pept. Sci."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Mazhar, Z., Akhtar, J., Alhodaib, A., Naz, T., Zafar, M.I., Iqbal, M.M., Fatima, H., and Naz, I. (2023). Efficacy of ZnO nanoparticles in Zn fortification and partitioning of wheat and rice grains under salt stress. Sci. Rep., 13.","DOI":"10.1038\/s41598-022-26039-8"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2968","DOI":"10.1039\/D0EN00582G","article-title":"Comparative physiological and metabolomics analysis reveals that single-walled carbon nanohorns and ZnO nanoparticles affect salt tolerance in Sophora alopecuroides","volume":"7","author":"Wan","year":"2020","journal-title":"Environ. Sci. Nano"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Shen, M., Liu, W., Zeb, A., Lian, J., Wu, J., and Lin, M. (2022). Bioaccumulation and phytotoxicity of ZnO nanoparticles in soil-grown Brassica chinensis L. and potential risks. J. Environ. Manag., 306.","DOI":"10.1016\/j.jenvman.2022.114454"},{"key":"ref_31","first-page":"62","article-title":"Effects of planting density on the growth, taproots yield and quality of Glycyrrhiza uralensis","volume":"46","author":"Jia","year":"2023","journal-title":"Legume Res."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Mousavi, S.S., Karami, A., Saharkhiz, M.J., Etemadi, M., and Ravanbakhsh, M. (2022). Microbial amelioration of salinity stress in endangered accessions of Iranian licorice (Glycyrrhiza glabra L.). BMC Plant Biol., 22.","DOI":"10.1186\/s12870-022-03703-9"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.plaphy.2013.11.001","article-title":"Nitric oxide-activated hydrogen sulfide is essential for cadmium stress response in bermudagrass (Cynodon dactylon (L). Pers.)","volume":"74","author":"Shi","year":"2014","journal-title":"Plant Physiol. Biochem."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1007\/s11099-013-0041-2","article-title":"Interaction of salicylic acid and ethylene and their effects on some physiological and biochemical parameters in canola plants (Brassica napus L.)","volume":"51","author":"Tirani","year":"2013","journal-title":"Photosynthetica"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1016\/j.jff.2014.12.003","article-title":"Chemical profile and inhibition of \u03b1-glycosidase and protein tyrosine phosphatase 1B (PTP1B) activities by flavonoids from licorice (Glycyrrhiza uralensis Fisch)","volume":"14","author":"Guo","year":"2015","journal-title":"J. Funct. Foods"},{"key":"ref_36","unstructured":"Feng, W., Wang, W.Q., and Zhao, P.R. (2007). Study on methods in determination of general flavonoids in Glycyrrhiza uralensis Fisch with ultraviolet spectrophotometry. J. Lishizhen Tradit. Chin. Med., 18."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Zhang, X., He, P., Guo, R., Huang, K., and Huang, X. (2023). Effects of salt stress on root morphology, carbon and nitrogen metabolism, and yield of Tartary buckwheat. Sci. Rep., 13.","DOI":"10.1038\/s41598-023-39634-0"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1111\/j.1469-8137.2005.01487.x","article-title":"Genes and salt tolerance: Bringing them together","volume":"167","author":"Munns","year":"2005","journal-title":"New Phytol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Ahmed, M., Marrez, D.A., Rizk, R., Zedan, M., Abdul-Hamid, D., Decsi, K., Kov\u00e1cs, G.P., and T\u00f3th, Z. (2024). The influence of Zinc oxide nanoparticles and salt stress on the morphological and some biochemical characteristics of Solanum lycopersicum L. plants. Plants, 13.","DOI":"10.3390\/plants13101418"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Dimkpa, C.O., Andrews, J., Sanabria, J., Bindraban, P.S., Singh, U., Elmer, W.H., Gardea-Torresdey, J.L., and White, J.C. (2020). Interactive effects of drought, organic fertilizer, and zinc oxide nanoscale and bulk particles on wheat performance and grain nutrient accumulation. Sci. Total Environ., 722.","DOI":"10.1016\/j.scitotenv.2020.137808"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Zhang, H., Zhou, Q., Liu, R., Zhao, Z., Liu, J., Siddique, K.H.M., and Mao, H. (2024). Enhancing zinc biofortification and mitigating cadmium toxicity in soil-earthworm-spinach systems using different zinc sources. J. Hazard. Mater., 476.","DOI":"10.1016\/j.jhazmat.2024.135243"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"11758","DOI":"10.1021\/jp301671b","article-title":"Dissolution kinetics and solubility of ZnO nanoparticles followed by AGNES","volume":"116","author":"David","year":"2012","journal-title":"J. Phys. Chem. C"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Elsherif, D.E., Abd-ElShafy, E., and Khalifa, A.M. (2023). Impacts of ZnO as a nanofertilizer on fenugreek: Some biochemical parameters and SCoT analysis. J. Genet. Eng. Biotechnol., 21.","DOI":"10.1186\/s43141-023-00501-0"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Francis, D.V., Sood, N., and Gokhale, T. (2022). Biogenic CuO and ZnO nanoparticles as nanofertilizers for sustainable growth of Amaranthus hybridus. Plants, 11.","DOI":"10.3390\/plants11202776"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Dang, K., Wang, Y., Tian, H., Bai, J., Cheng, X., Guo, L., Zhang, Q., Geng, Y., and Shao, X. (2024). Impact of ZnO NPs on photosynthesis in rice leaves plants grown in saline-sodic soil. Sci. Rep., 14.","DOI":"10.1038\/s41598-024-66935-9"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Ahmed, M., Decsi, K., and T\u00f3th, Z. (2022). Different tactics of synthesized Zinc oxide nanoparticles, homeostasis ions, and phytohormones as regulators and adaptatively parameters to alleviate the adverse effects of salinity stress on plants. Life, 13.","DOI":"10.3390\/life13010073"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., Raihan, M.R.H., Masud, A.A.C., Rahman, K., Nowroz, F., Rahman, M., Nahar, K., and Fujita, M. (2021). Regulation of reactive oxygen species and antioxidant defense in plants under salinity. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22179326"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1046\/j.1469-8137.2000.00630.x","article-title":"Tansley Review No. 111: Possible roles of zinc in protecting plant cells from damage by reactive oxygen species","volume":"146","author":"Cakmak","year":"2000","journal-title":"New Phytol."},{"key":"ref_49","first-page":"1351","article-title":"Plant responses to abiotic stresses: Heavy metal-induced oxidative stress and protection by mycorrhization","volume":"53","author":"Polle","year":"2002","journal-title":"J. Exp. Bot."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.molp.2016.09.001","article-title":"Plant carbonic anhydrases: Structures, locations, evolution, and physiological roles","volume":"10","author":"DiMario","year":"2017","journal-title":"Mol. Plant"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Gogna, M., and Bhatla, S.C. (2019). Biochemical mechanisms regulating salt tolerance in sunflower. Plant Signal. Behav., 14.","DOI":"10.1080\/15592324.2019.1670597"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Pallavi, S., Bhushan, J.A., Shanker, D.R., and Mohammad, P. (2012). Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. J. Bot., 2012.","DOI":"10.1155\/2012\/217037"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Hussan, M.U., Hussain, S., Hafeez, M.B., Ahmed, S., Hassan, M.U., Jabeen, S., Yan, M., and Wang, Q. (2024). Comparative role of calcium oxide nanoparticles and calcium bulk fertilizer to alleviate cadmium toxicity by modulating oxidative stress, photosynthetic performance and antioxidant-defense genes expression in alfalfa. Plant Physiol. Biochem., 215.","DOI":"10.1016\/j.plaphy.2024.109002"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Mirakhorli, T., Ardebili, Z.O., Ladan-Moghadam, A., and Danaee, E. (2021). Bulk and nanoparticles of zinc oxide exerted their beneficial effects by conferring modifications in transcription factors, histone deacetylase, carbon and nitrogen assimilation, antioxidant biomarkers, and secondary metabolism in soybean. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0256905"},{"key":"ref_55","first-page":"422","article-title":"Exogenous silicium and zinc increase antioxidant enzyme activity and alleviate salt stress in leaves of sunflower","volume":"9","author":"Ebrahimian","year":"2011","journal-title":"J. Food Agric. Environ."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Kaur, S., Garg, T., Joshi, A., Awasthi, A., Kumar, V., and Kumar, A. (2024). Potential effects of metal oxide nanoparticles on leguminous plants: Practical implications and future perspectives. Sci. Hortic., 331.","DOI":"10.1016\/j.scienta.2024.113146"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1321","DOI":"10.1016\/j.plantsci.2004.01.011","article-title":"Zinc protects chloroplasts and associated photochemical functions in cadmium exposed Ceratophyllum demersum L., a freshwater macrophyte","volume":"166","author":"Aravind","year":"2004","journal-title":"Plant Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"3132","DOI":"10.1016\/j.sjbs.2020.08.005","article-title":"The potential mitigation effect of ZnO nanoparticles on Abelmoschus esculentus L. Moench metabolism under salt stress conditions","volume":"27","author":"Alabdallah","year":"2020","journal-title":"Saudi J. Biol. Sci."},{"key":"ref_59","first-page":"242","article-title":"Zinc metalloenzymes in plants","volume":"43","author":"Castillo","year":"2018","journal-title":"Interciencia"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1080\/17429145.2017.1385867","article-title":"Zinc application mitigates the adverse effects of NaCl stress on mustard [Brassica juncea (L.) Czern & Coss] through modulating compatible organic solutes, antioxidant enzymes, and flavonoid content","volume":"12","author":"Ahmad","year":"2017","journal-title":"J. Plant Interact."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.sajb.2019.12.014","article-title":"Physiological and molecular mechanisms of salinity tolerance in grafted cucumber","volume":"130","author":"Elsheery","year":"2020","journal-title":"S. Afr. J. Bot."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1080\/03650340802393303","article-title":"Genotypic differences in plant growth, osmotic and antioxidative defence of Cajanus cajan (L.) Millsp. modulated by salt stress","volume":"55","author":"Garg","year":"2009","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/j.plaphy.2017.06.020","article-title":"Effects, tolerance mechanisms and management of salt stress in grain legumes","volume":"118","author":"Farooq","year":"2017","journal-title":"Plant Physiol. Biochem."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Mushtaq, N.U., Alghamdi, K.M., Saleem, S., Tahir, I., Bahieldin, A., Henrissat, B., Alghamdi, M.K., Rehman, R.U., and Hakeem, K.R. (2023). Exogenous zinc mitigates salinity stress by stimulating proline metabolism in proso millet (Panicum miliaceum L.). Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1053869"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"19989","DOI":"10.1007\/s11356-016-7170-7","article-title":"Seed priming with polyethylene glycol induces antioxidative defense and metabolic regulation of rice under nano-ZnO stress","volume":"23","author":"Sheteiwy","year":"2016","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"4112","DOI":"10.1105\/tpc.110.082685","article-title":"Triterpene functional genomics in licorice for identification of CYP72A154 involved in the biosynthesis of glycyrrhizin","volume":"23","author":"Seki","year":"2011","journal-title":"Plant Cell"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Mousavi, S.S., Karami, A., Saharkhiz, M.J., Etemadi, M., and Zarshenas, M.M. (2022). Evaluation of metabolites in Iranian Licorice accessions under salinity stress and Azotobacter sp. Inoculation. Sci. Rep., 12.","DOI":"10.1038\/s41598-022-20366-6"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Takla, S.S., Shawky, E., Mahgoub, Y.A., and Darwish, R.S. (2023). Tracking the effect of roasting and fermentation on the metabolites of licorice root (Glycyrrhiza glabra L.) using UPLC-MS analysis combined with multivariate statistical analysis. BMC Complement. Med. Ther., 23.","DOI":"10.1186\/s12906-023-04239-7"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1515\/biol-2017-0016","article-title":"The influence of soil salinity on volatile organic compounds emission and photosynthetic parameters of Solanum lycopersicum L. varieties","volume":"12","author":"Tomescu","year":"2017","journal-title":"Open Life Sci."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1480","DOI":"10.1021\/acs.jafc.9b04887","article-title":"Comparative proteomic analysis reveals the molecular mechanisms underlying the accumulation difference of bioactive constituents in Glycyrrhiza uralensis Fisch under salt stress","volume":"68","author":"Wang","year":"2020","journal-title":"J. Agric. Food Chem."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1080\/13880209.2016.1225775","article-title":"The anti-inflammatory activity of licorice, a widely used Chinese herb","volume":"55","author":"Yang","year":"2017","journal-title":"Pharm. Biol."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Pradeep, M., Saxena, M., Mondal, D., and Franklin, G. (2024). Do nanoparticles delivered to roots affect plant secondary metabolism?. A comprehensive analysis in float seedling cultures of Hypericum perforatum L. Chemosphere, 356.","DOI":"10.1016\/j.chemosphere.2024.141789"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Naseem, J., Shah, A.A., Usman, S., Ahmed, S., Gatasheh, M.K., Shaffique, S., and Javad, S. (2025). Green synthesized FeNPs ameliorate drought stress in Spinacia oleracea L. through improved photosynthetic capacity, redox balance, and antioxidant defense. Sci. Rep., 15.","DOI":"10.1038\/s41598-024-84061-4"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Singh, P., Arif, Y., Siddiqui, H., Sami, F., Zaidi, R., Azam, A., Alam, P., and Hayat, S. (2021). Nanoparticles enhances the salinity toxicity tolerance in Linum usitatissimum L. by modulating the antioxidative enzymes, photosynthetic efficiency, redox status and cellular damage. Ecotoxicol. Environ. Saf., 213.","DOI":"10.1016\/j.ecoenv.2021.112020"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Shibli, R., Mohusaien, R., AbuZurayk, R., Qudah, T., and Tahtamouni, R. (2022). Silver nanoparticles (Ag NPs) boost mitigation powers of Chenopodium quinoa (Q6 line) grown under in vitro salt-stressing conditions. Water, 14.","DOI":"10.3390\/w14193099"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Jangid, H., and Kumar, G. (2025). Ecotoxicity of fungal-synthesized silver nanoparticles: Mechanisms, impacts, and sustainable mitigation strategies. 3 Biotech, 15.","DOI":"10.1007\/s13205-025-04266-w"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/j.apsoil.2018.01.007","article-title":"Engineered nanoparticles effects in soil-plant system: Basil (Ocimum basilicum L.) study case","volume":"123","author":"Antisari","year":"2018","journal-title":"Appl. Soil Ecol."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Gurusamy, M., Sellavel, M., and Kuppuvelsamy, V.B. (2024). A sustainable green synthesis for photocatalytic and antibacterial activity of zinc oxide nanoparticles using Cucumis maderaspatanus leaf extract. Desalin. Water Treat., 319.","DOI":"10.1016\/j.dwt.2024.100457"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Ramaprabha, K., and Kumar, S.V. (2025). Effective photocatalytic degradation and kinetic modelling of azo dyes by zinc oxide nanoparticles from Brevibacterium casei. Desalin. Water Treat., 321.","DOI":"10.1016\/j.dwt.2024.100936"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"3391","DOI":"10.1039\/C4TB00069B","article-title":"Preparation and characterization of novel water-based biodegradable polyurethane nanoparticles encapsulating superparamagnetic iron oxide and hydrophobic drugs","volume":"2","author":"Chen","year":"2014","journal-title":"J. Mater. Chem. B"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"478","DOI":"10.1016\/j.fct.2017.09.037","article-title":"Perspectives from the NanoSafety Modelling Cluster on the validation criteria for (Q)SAR models used in nanotechnology","volume":"112","author":"Puzyn","year":"2018","journal-title":"Food Chem. Toxicol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/12\/1763\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:49:00Z","timestamp":1760032140000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/12\/1763"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,6,9]]},"references-count":81,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2025,6]]}},"alternative-id":["plants14121763"],"URL":"https:\/\/doi.org\/10.3390\/plants14121763","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,6,9]]}}}