{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T14:39:57Z","timestamp":1792593597829,"version":"4.5.0"},"reference-count":81,"publisher":"American Chemical Society (ACS)","issue":"41","license":[{"start":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T00:00:00Z","timestamp":1791590400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100003825","name":"Magyar Tudom??nyos Akad??mia","doi-asserted-by":"publisher","award":["LENDULET_2023-40"],"award-info":[{"award-number":["LENDULET_2023-40"]}],"id":[{"id":"10.13039\/501100003825","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["K 135303"],"award-info":[{"award-number":["K 135303"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018818","name":"National Research, Development and Innovation Office","doi-asserted-by":"publisher","award":["K 146733"],"award-info":[{"award-number":["K 146733"]}],"id":[{"id":"10.13039\/501100018818","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,21]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Zinc (Zn) deficiency severely limits plant growth and yield, highlighting the need for strategies to improve Zn nutrition. Here, Zn oxide nanoparticles (nZnOs) of two sizes (\u223c8 nm and \u223c45 nm, 100 mg\u00b7L\u20131) were evaluated as seed priming agents in tomato grown under Zn-deficient conditions. Both nanoparticle sizes were internalized into embryonic tissues and showed distinct intracellular localization. While 45 nm nZnO and bulk Zn disturbed seed nutrient homeostasis, 8 nm nZnO increased seed Zn content more efficiently, with milder effects on other nutrients. Across developmental stages, nZnO priming improved Zn supply and nutrient distribution more effectively than bulk Zn. At the vegetative stage, 45 nm nZnO enhanced overall biomass, whereas 8 nm nZnO promoted root-specific growth under Zn deficiency. nZnO priming increased root Zn accumulation, particularly with 8 nm nZnO, and alleviated Zn deficiency-induced nonspecific metal accumulation. Zn deficiency-induced disturbances in redox and nitrosative balance were reversed by nZnO priming, accompanied by reduced RNA nitration and a shift toward a catalase-dominated hydrogen peroxide detoxification system. Both nanoparticle sizes exhibited peroxidase-like activity in cell-free assays, indicating nanozyme potential; however, this activity did not translate directly to the in planta responses. Under Zn deficiency, nZnO priming suppressed stress-related metabolic pathways in leaves, while enhancing amino acid, polyamine, and structural carbohydrate metabolism in roots. Zn deficiency-induced transcriptional repression, particularly in leaves, was partially restored, with affected genes enriched in stress-, phytohormone-, and detoxification-related pathways. Overall, size-dependent nZnO seed priming alleviates Zn deficiency in tomato. 45 nm nZnO mainly enhances biomass through improved nutrient homeostasis and redox regulation, while both sizes contribute to restored Zn availability and antioxidant reprogramming under Zn deficiency.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c02493","type":"journal-article","created":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T15:01:57Z","timestamp":1791644517000},"page":"32067-32086","source":"Crossref","is-referenced-by-count":0,"title":["Size-Dependent\nZinc Delivery and Nanozyme-Mediated\nStress Alleviation by ZnO Nanoparticles in Zinc-Deficient Tomato"],"prefix":"10.1021","volume":"74","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4979-5986","authenticated-orcid":true,"given":"Selahattin","family":"Kondak","sequence":"first","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Do\u0301ra","family":"Kondak","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tama\u0301s","family":"Bodor","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]},{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ga\u0301bor","family":"Fejes","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]},{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Attila","family":"O\u0308rdo\u0308g","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7054-0975","authenticated-orcid":true,"given":"Andrea","family":"Ro\u0301nava\u0301ri","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6720"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9406-8596","authenticated-orcid":true,"given":"Zolta\u0301n","family":"Ko\u0301nya","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6720"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Re\u0301ka","family":"Szo\u030bllo\u030bsi","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andra\u0301s","family":"Kukri","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Attila","family":"Gyenesei","sequence":"additional","affiliation":[{"name":"University of Pe\u0301cs , , ,","place":["Pe\u0301cs, Hungary, 7624"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pe\u0301ter","family":"Urba\u0301n","sequence":"additional","affiliation":[{"name":"University of Pe\u0301cs , , ,","place":["Pe\u0301cs, Hungary, 7624"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jo\u0301zsef","family":"Kun","sequence":"additional","affiliation":[{"name":"University of Pe\u0301cs , , ,","place":["Pe\u0301cs, Hungary, 7624"]},{"name":"Medical School University of Pe\u0301cs , , ,","place":["Pe\u0301cs, Hungary, 7624"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Gracheva","sequence":"additional","affiliation":[{"name":"HUN-REN Centre for Energy Research , , ,","place":["Budapest, Hungary, 1121"]},{"name":"ELTE Eo\u0308tvo\u0308s Lora\u0301nd University , , ,","place":["Budapest, Hungary, 1117"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"A\u0301da\u0301m","family":"Solti","sequence":"additional","affiliation":[{"name":"ELTE Eo\u0308tvo\u0308s Lora\u0301nd University , , ,","place":["Budapest, Hungary, 1117"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kinga","family":"Benczu\u0301r","sequence":"additional","affiliation":[{"name":"HUN-REN Centre for Agricultural Research , , ,","place":["Martonva\u0301sa\u0301r, Hungary, 2462"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3468-962X","authenticated-orcid":true,"given":"Magda","family":"Pa\u0301l","sequence":"additional","affiliation":[{"name":"HUN-REN Centre for Agricultural Research , , ,","place":["Martonva\u0301sa\u0301r, Hungary, 2462"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabriella","family":"Szalai","sequence":"additional","affiliation":[{"name":"HUN-REN Centre for Agricultural Research , , ,","place":["Martonva\u0301sa\u0301r, Hungary, 2462"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ildiko\u0301","family":"Domonkos","sequence":"additional","affiliation":[{"name":"HUN-REN Biological Research Centre , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Titanilla","family":"Szo\u0308gi","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6720"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fernando A.","family":"Casian Plaza","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6720"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1799-5329","authenticated-orcid":true,"given":"Ga\u0301bor","family":"Galba\u0301cs","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6720"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7819-4672","authenticated-orcid":true,"given":"Zsuzsanna","family":"Kolbert","sequence":"additional","affiliation":[{"name":"University of Szeged , , ,","place":["Szeged, Hungary, 6726"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,10,10]]},"reference":[{"issue":"2","key":"2026102109082079800_cit1","doi-asserted-by":"publisher","first-page":"112","DOI":"10.33545\/2664844X.2024.v6.i2b.213","article-title":"Understanding the vital role of Zn in plant nutrition","volume":"6","author":"Sasmal","year":"2024","journal-title":"Int. J. Agric. Food Sci."},{"key":"2026102109082079800_cit2","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1016\/j.plantsci.2014.07.010","article-title":"Role of\nGSH homeostasis under Zn toxicity in plants with different Zn tolerance","volume":"227","author":"Barrameda-Medina","year":"2014","journal-title":"Plant Sci."},{"issue":"9","key":"2026102109082079800_cit3","doi-asserted-by":"publisher","first-page":"396","DOI":"10.3390\/agriculture10090396","article-title":"The Critical\nRole of Zinc in Plants Facing the Drought Stress","volume":"10","author":"Umair Hassan","year":"2020","journal-title":"Agriculture"},{"issue":"3","key":"2026102109082079800_cit4","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1007\/s10725-025-01294-7","article-title":"Enhancing soil health\nand crop productivity: the role of zinc-solubilizing bacteria in sustainable\nagriculture","volume":"105","author":"Sethi","year":"2025","journal-title":"Plant Growth Regul."},{"key":"2026102109082079800_cit5","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/B978-0-323-91314-0.00011-9","article-title":"Zn deficiency in plants: an\ninsight into fortification strategies","volume-title":"Zn in Plants","author":"Sharma","year":"2025"},{"key":"2026102109082079800_cit6","doi-asserted-by":"publisher","first-page":"153419","DOI":"10.1016\/j.jplph.2021.153419","article-title":"Molecular regulation\nof Zn deficiency responses in plants","volume":"261","author":"Zeng","year":"2021","journal-title":"J. Plant\nPhysiol."},{"key":"2026102109082079800_cit7","doi-asserted-by":"publisher","first-page":"105330","DOI":"10.1016\/j.envexpbot.2023.105330","article-title":"Revealing\nthe molecular mechanisms of Zn accumulation and Zn deficiency responses\nin quinoa, analyzed by high-throughput gene expression profiling under\nZn depletion and resupply","volume":"210","author":"Lv","year":"2023","journal-title":"Environ. Exp. Bot."},{"key":"2026102109082079800_cit8","doi-asserted-by":"publisher","first-page":"1597149","DOI":"10.3389\/fagro.2025.1597149","article-title":"Microbe-mediated regulation\nin Zn-contaminated soils: the synergistic role of hyperaccumulator\nplants and Zn-tolerant rhizobacteria","volume":"7","author":"Rani","year":"2025","journal-title":"Front.\nAgron."},{"issue":"1","key":"2026102109082079800_cit9","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1111\/ejss.12437","article-title":"Agronomic biofortification\nof cereals with Zn: a review","volume":"69","author":"C\u0327akmak","year":"2018","journal-title":"Eur. J. Soil Sci."},{"issue":"26","key":"2026102109082079800_cit10","doi-asserted-by":"publisher","first-page":"6462","DOI":"10.1021\/acs.jafc.7b02150","article-title":"Nanofertilizers:\nnew products for the industry?","volume":"66","author":"Dimkpa","year":"2018","journal-title":"J. Agric. Food\nChem."},{"issue":"8","key":"2026102109082079800_cit11","doi-asserted-by":"publisher","first-page":"1646","DOI":"10.3390\/agronomy14081646","article-title":"Fertilizers\nbased\non nanoparticles as sources of macro-and microelements for plant crop\ngrowth: A review","volume":"14","author":"Semenova","year":"2024","journal-title":"Agronomy"},{"key":"2026102109082079800_cit12","doi-asserted-by":"publisher","first-page":"64","DOI":"10.3389\/fchem.2015.00064","article-title":"Nanopesticides\nand nanofertilizers: emerging contaminants or opportunities for risk\nmitigation?","volume":"3","author":"Kah","year":"2015","journal-title":"Front. Chem."},{"issue":"2","key":"2026102109082079800_cit13","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1007\/s11051-015-2907-7","article-title":"A review of the use of engineered nanomaterials to\nsuppress plant disease and enhance crop yield","volume":"17","author":"Servin","year":"2015","journal-title":"J. Nanopart. Res."},{"issue":"15","key":"2026102109082079800_cit14","doi-asserted-by":"publisher","first-page":"2430","DOI":"10.3390\/plants14152430","article-title":"ZnO Nanoparticles:\nAdvancing Agricultural Sustainability","volume":"14","author":"Ravishankar","year":"2025","journal-title":"Plants"},{"issue":"2","key":"2026102109082079800_cit15","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1016\/j.aoas.2020.08.001","article-title":"Accumulation\nof nanoparticles in\nthe soil-plant systems and their effects on human health","volume":"65","author":"Rajput","year":"2020","journal-title":"Ann. Agric. Sci."},{"key":"2026102109082079800_cit16","doi-asserted-by":"publisher","first-page":"148927","DOI":"10.1016\/j.scitotenv.2021.148927","article-title":"Interaction\nof different-sized ZnO nanoparticles with maize (Zea mays): Accumulation,\nbiotransformation and phytotoxicity","volume":"796","author":"Lv","year":"2021","journal-title":"Sci. Total\nEnviron."},{"key":"2026102109082079800_cit17","doi-asserted-by":"publisher","first-page":"78","DOI":"10.3389\/fchem.2017.00078","article-title":"Impact of\nmetal and metal oxide nanoparticles\non plant: a critical review","volume":"5","author":"Rastogi","year":"2017","journal-title":"Front. Chem."},{"issue":"1","key":"2026102109082079800_cit18","doi-asserted-by":"publisher","first-page":"8263","DOI":"10.1038\/s41598-017-08669-5","article-title":"Nanopriming\ntechnology for enhancing germination and starch metabolism of aged\nrice seeds using phytosynthesized silver nanoparticles","volume":"7","author":"Mahakham","year":"2017","journal-title":"Sci. Rep."},{"issue":"1","key":"2026102109082079800_cit19","doi-asserted-by":"publisher","first-page":"5037","DOI":"10.1038\/s41598-020-61696-7","article-title":"Nanoparticle-mediated\nseed priming improves germination, growth, yield, and quality of watermelons\n(Citrullus lanatus) at multi-locations in Texas","volume":"10","author":"Acharya","year":"2020","journal-title":"Sci. Rep."},{"key":"2026102109082079800_cit20","volume-title":"Plant Exposure to Engineered\nNanoparticles: uptake, Transformation, Molecular and Physiological\nResponses.","author":"Rico","year":"2022"},{"key":"2026102109082079800_cit21","doi-asserted-by":"publisher","first-page":"100124","DOI":"10.1016\/j.plana.2024.100124","article-title":"Harnessing\nnanotechnology for sustainable\nagriculture: from seed priming to encapsulation","volume":"11","author":"Mahra","year":"2025","journal-title":"Plant Nano Biol."},{"key":"2026102109082079800_cit22","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1016\/j.plaphy.2016.07.030","article-title":"An overview on manufactured\nnanoparticles in plants: uptake, translocation, accumulation and phytotoxicity","volume":"110","author":"Tripathi","year":"2017","journal-title":"Plant Physiol. Biochem."},{"key":"2026102109082079800_cit23","doi-asserted-by":"publisher","first-page":"1123931","DOI":"10.3389\/fsoil.2023.1123931","article-title":"Fertilizers\nfor food and nutrition\nsecurity in sub-Saharan Africa: an overview of soil health implications","volume":"3","author":"Dimkpa","year":"2023","journal-title":"Front. Soil Sci."},{"key":"2026102109082079800_cit24","doi-asserted-by":"publisher","first-page":"126419","DOI":"10.1016\/j.chemosphere.2020.126419","article-title":"Nitro-oxidative\nsignalling induced by chemically synthetized zinc oxide nanoparticles\n(ZnO NPs) in Brassica species","volume":"251","author":"Molna\u0301r","year":"2020","journal-title":"Chemosphere"},{"key":"2026102109082079800_cit25","doi-asserted-by":"publisher","first-page":"111158","DOI":"10.1016\/j.ecoenv.2020.111158","article-title":"ZnO nanoparticles induce cell wall\nremodeling and modify\nROS\/RNS signalling in roots of Brassica seedlings","volume":"206","author":"Molna\u0301r","year":"2020","journal-title":"Ecotoxicol. Environ. Saf."},{"issue":"10","key":"2026102109082079800_cit26","doi-asserted-by":"publisher","first-page":"6059","DOI":"10.1021\/la200570n","article-title":"Aggregation and\ndissolution of 4 nm ZnO nanoparticles in aqueous environments: influence\nof pH, ionic strength, size, and adsorption of humic acid","volume":"27","author":"Bian","year":"2011","journal-title":"Langmuir"},{"issue":"8","key":"2026102109082079800_cit27","doi-asserted-by":"publisher","first-page":"9803","DOI":"10.1016\/j.ceramint.2013.04.110","article-title":"Synthesis,\ncharacterization and application\nof Zn oxide nanoparticles (n-ZnO)","volume":"39","author":"Srivastava","year":"2013","journal-title":"Ceram. Int."},{"key":"2026102109082079800_cit28","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1515\/abcsb-2015-0008","article-title":"The alleviation of the adverse effects of salt stress\nin the tomato plant by salicylic acid shows a time-and organ-specific\nantioxidant response","volume":"57","author":"Tari","year":"2015","journal-title":"Acta Biol. Cracov., Ser.\nBot."},{"issue":"1","key":"2026102109082079800_cit29","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1093\/jxb\/32.1.93","article-title":"Leaf senescence: correlated with\nincreased levels of membrane permeability and lipid peroxidation,\nand decreased levels of superoxide dismutase and catalase","volume":"32","author":"Dhindsa","year":"1981","journal-title":"J. Exp. Bot."},{"issue":"5","key":"2026102109082079800_cit30","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/S0176-1617(85)80081-X","article-title":"Effect\nof paclobutrazol on the activities of some enzymes of activated oxygen\nmetabolism and lipid peroxidation in senescing soybean leaves","volume":"121","author":"Upadhyaya","year":"1985","journal-title":"J. Plant Physiol."},{"issue":"1\u20132","key":"2026102109082079800_cit31","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A rapid\nand sensitive method for the quantitation of microgram quantities\nof protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."},{"issue":"8","key":"2026102109082079800_cit32","doi-asserted-by":"publisher","first-page":"2053","DOI":"10.3390\/agronomy13082053","article-title":"Contribution\nof glutathione transferases in the selective and light-dependent effect\nof flumioxazin on winter wheat (Triticum aestivum L.) and its typical weed common poppy (Papaver rhoeas L.)","volume":"13","author":"Galle\u0301","year":"2023","journal-title":"Agronomy"},{"issue":"2","key":"2026102109082079800_cit33","doi-asserted-by":"publisher","first-page":"913","DOI":"10.1039\/D3NR04336C","article-title":"Facile green synthesis of wasted hop-based zinc oxide nanozymes as\nperoxidase-like catalysts for colorimetric analysis","volume":"16","author":"Liu","year":"2024","journal-title":"Nanoscale"},{"issue":"1","key":"2026102109082079800_cit34","first-page":"37","article-title":"In vivo and\nin vitro studies on fluorophore-specificity","volume":"56","author":"Kolbert","year":"2012","journal-title":"Acta Biol. Szeged."},{"issue":"3","key":"2026102109082079800_cit35","doi-asserted-by":"publisher","first-page":"1158","DOI":"10.1104\/pp.108.122598","article-title":"The level of free intracellular Zn\nmediates programmed cell death\/cell survival decisions in plant embryos","volume":"147","author":"Helmersson","year":"2008","journal-title":"Plant Physiol."},{"issue":"15","key":"2026102109082079800_cit36","doi-asserted-by":"publisher","first-page":"5677","DOI":"10.1093\/jxb\/ers222","article-title":"Selenite-induced hormonal\nand signalling mechanisms during root growth of Arabidopsis thaliana\nL","volume":"63","author":"Lehotai","year":"2012","journal-title":"J. Exp. Bot."},{"issue":"2","key":"2026102109082079800_cit37","doi-asserted-by":"publisher","first-page":"147","DOI":"10.29244\/jtcs.11.02.147-154","article-title":"Modification of the spectrophotometric\nanalysis protocol by sims d. and gamon to analyze leaf pigment content\nusing green spinach (Amaranthus hybridus) as a model\nplant","volume":"11","author":"Yudiansyah","year":"2024","journal-title":"J. Trop. Crop Sci."},{"issue":"2\u20133","key":"2026102109082079800_cit38","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1016\/S0034-4257(02)00010-X","article-title":"Relationships between\nleaf pigment content and spectral\nreflectance across a wide range of species, leaf structures and developmental\nstages","volume":"81","author":"Sims","year":"2002","journal-title":"Remote Sens. 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