{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T05:04:50Z","timestamp":1784178290461,"version":"3.55.0"},"reference-count":67,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2026,7,11]],"date-time":"2026-07-11T00:00:00Z","timestamp":1783728000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Iron plays a key role in plant chemical processes and photosynthesis. The aim of this study was to investigate the effect of various iron concentrations on total phenolic compound levels and photosynthetic efficiency in Melissa officinalis. It is a very popular herb, widely cultivated worldwide for its medicinal properties. Herbs were grown under controlled laboratory conditions. Gas exchange parameters, chlorophyll index, and total content of phenolic compounds were determined. Additionally, iron, manganese, copper, and zinc concentrations in either soil or plant tissues were analyzed. The hypothesis that iron supplementation significantly affects photosynthesis and total phenolic compounds in lemon balm was confirmed by one-way analysis of variance. In particular, the beneficial biofortification at 100 and 200 \u00b5g\/g iron doses generates a moderate increase in phenolic compounds and simultaneously does not hamper plant growth and photosynthetic efficiency. However, iron doses above 500 \u00b5g\/g induce plant stress and trigger defense mechanisms that significantly increase phenolic compound production. This is accompanied by a decrease in biomass and photosynthetic efficiency.<\/jats:p>","DOI":"10.3390\/ijms27146211","type":"journal-article","created":{"date-parts":[[2026,7,13]],"date-time":"2026-07-13T08:53:58Z","timestamp":1783932838000},"page":"6211","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Iron Nitrates Stimulate Phenolic Compound Production by Melissa officinalis L."],"prefix":"10.3390","volume":"27","author":[{"given":"Dorota","family":"Adamczyk-Szabela","sequence":"first","affiliation":[{"name":"Institute of General and Ecological Chemistry, Lodz University of Technology, Zeromskiego 116, 90-924 Lodz, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wojciech M.","family":"Wolf","sequence":"additional","affiliation":[{"name":"Institute of General and Ecological Chemistry, Lodz University of Technology, Zeromskiego 116, 90-924 Lodz, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-3385-9282","authenticated-orcid":false,"given":"Dawid","family":"Krakowiak","sequence":"additional","affiliation":[{"name":"Institute of General and Ecological Chemistry, Lodz University of Technology, Zeromskiego 116, 90-924 Lodz, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,11]]},"reference":[{"key":"ref_1","first-page":"55","article-title":"Effect of organic cultivation on yield and quality of lemon balm herb (Melissa officinalis L.)","volume":"14","author":"Mordalski","year":"2015","journal-title":"Acta Sci. Pol. Hortorum Cultus"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"e70864","DOI":"10.1002\/fsn3.70864","article-title":"Bioactive Compounds, Medicinal Benefits, and Contemporary Extraction Methods for Lemon Balm (Melissa officinalis)","volume":"13","author":"Awlqadr","year":"2025","journal-title":"Food Sci. Nutr."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1039\/C7MT00136C","article-title":"Iron homeostasis in plants\u2014A brief overview","volume":"9","author":"Connorton","year":"2017","journal-title":"Metallomics"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1016\/j.febslet.2007.04.043","article-title":"Mining iron: Iron uptake and transport in plants","volume":"581","author":"Kim","year":"2007","journal-title":"FEBS Lett."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"102927","DOI":"10.1016\/j.redox.2023.102927","article-title":"Polyphenols and the regulation of NADPH oxidases","volume":"67","author":"Fragaa","year":"2023","journal-title":"Red. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Yahfoufi, N., Alsadi, N., Jambi, M., and Matar, C. (2018). The immunomodulatory and anti-inflammatory role of polyphenols nutrients. Nutrients, 10.","DOI":"10.3390\/nu10111618"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1016\/S1369-5266(99)80043-0","article-title":"Iron acquisition by plants","volume":"2","author":"Mori","year":"1999","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Kabata-Pendias, A., and Pendias, H. (2001). Trace Elements in Soils and Plants, CRC Press. [3rd ed.].","DOI":"10.1201\/9781420039900"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Velez-Bermudez, I.C., and Schmidt, W. (2023). Iron sensing in plants. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1145510"},{"key":"ref_10","unstructured":"Korzeniowska, J., Stanis\u0142awska-Glubiak, E., Jadczyszyn, T., and Lipi\u0144ski, W. (2021). Fertilizing Agricultural Crops with Microelements, National Centre for Research and Development, IUNG-PIB."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/BF00011851","article-title":"Solubility and dissolution of iron oxides","volume":"130","author":"Schwertmann","year":"1991","journal-title":"Plant Soil"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1007\/BF00011852","article-title":"Iron oxide solubilization by organic matter and its effect in iron availability","volume":"130","author":"Lindsay","year":"1991","journal-title":"Plant Soil"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3583","DOI":"10.1021\/acs.jafc.2c07727","article-title":"Biofortification of crops to fight anemia: Role of vacuolar iron trans-porters","volume":"71","author":"Krishna","year":"2023","journal-title":"J. Agric. Food Chem."},{"key":"ref_14","first-page":"11","article-title":"Iron transport and its regulation in plants","volume":"133","author":"Takanori","year":"2018","journal-title":"Free Radic. Biol. Med."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1","DOI":"10.7831\/ras.3.1","article-title":"Role of iron in plant growth and metabolism","volume":"3","author":"Rout","year":"2015","journal-title":"Rev. Agric. Sci."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Singh, V.P., Singh, S., Tripathi, D.K., Prasad, S.M., and Chauhan, D.K. (2021). Metal-based nanoparticles\u2019 interactions with plants. Plant Responses to Nanomaterials Recent Interventions and Physiological and Biochemical Responses, Springer.","DOI":"10.1007\/978-3-030-36740-4"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Ning, X., Lin, M., Huang, G., Mao, J., Gao, Z., and Wang, X. (2023). Research progress on iron absorption, transport, and molecular regulation strategy in plants. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1190768"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Johnson, D.B., Kanao, T., and Hedrich, S. (2012). Redox transformations of iron at extremely low pH: Fundamental and applied aspects. Front. Microbiol., 3.","DOI":"10.3389\/fmicb.2012.00096"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2154","DOI":"10.1093\/jxb\/erab012","article-title":"Using natural variation to understand plant responses to iron availability","volume":"72","author":"Miller","year":"2021","journal-title":"J. Exp. Bot."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1111\/j.1365-313X.2005.02624.x","article-title":"Rice plants take up iron as an Fe3+ phytosiderophore and as Fe2+","volume":"45","author":"Ishimaru","year":"2006","journal-title":"Plant J."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4","DOI":"10.36349\/easjpp.2024.v06i04.006","article-title":"Iron toxicity in plants: A Review","volume":"6","author":"Hamzah","year":"2024","journal-title":"EAS J. Pharm. Pharmacol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Aung, M.S., and Masuda, H. (2020). How does rice defend against excess iron?: Physiological and molecular mechanisms. Front. Plant Sci., 11, Correction in Front. Plant Sci. 2020, 11, 601527. https:\/\/doi.org\/10.3389\/fpls.2020.601527.","DOI":"10.3389\/fpls.2020.01102"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1104\/pp.106.092106","article-title":"Degradation of oxidized proteins by autophagy during oxidative stress in Arabidopsis","volume":"143","author":"Xiong","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s10681-008-9870-7","article-title":"QTL mapping for biomass and physiological parameters linked to resistance mechanisms to ferrous iron toxicity in rice","volume":"167","author":"Dufey","year":"2009","journal-title":"Euphytica"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.plantsci.2012.12.003","article-title":"Iron excess affects rice photosynthesis through stomatal and non-stomatal limitations","volume":"201","author":"Pereira","year":"2013","journal-title":"Plant Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1104\/pp.107.097162","article-title":"The FRD3-mediated efflux of citrate into the root vasculature is necessary for efficient iron translocation1[OA]","volume":"144","author":"Durrett","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_27","first-page":"1","article-title":"Iron deficiency in blackgram (Vigna mungo L.), Redox status and antioxidant activity","volume":"156","author":"Dey","year":"2020","journal-title":"Plant Biosyst."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2045","DOI":"10.1093\/jxb\/erab003","article-title":"All together now: Regulation of the iron deficiency response","volume":"72","author":"Riaz","year":"2021","journal-title":"J. Exp. Bot."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Schmidt, W., Thomine, S., and Buckhout, T.J. (2020). Editorial: Iron nutrition and interactions in plants. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01670"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1104\/pp.107.095794","article-title":"Iron deficiency-induced secretion of phenolics facilitates the reutilization of root apoplastic iron in red clover","volume":"44","author":"Jin","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"60","DOI":"10.4161\/psb.3.1.4902","article-title":"Iron deficiency-induced phenolics secretion plays multiple important roles in the plant iron acquisition underground","volume":"3","author":"Jin","year":"2008","journal-title":"Plant Signal. Behav."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Li, Y., Yan, X., Xu, Y., Pope, R., Spector, T.D., Falchi, M., Steves, C.J., Bell, J.T., Small, K.S., and Menni, C. (2025). Higher adherence to (poly)phenol-rich diet is associated with lower CVD risk in the Twins UK cohort. BMC Med., 23.","DOI":"10.1186\/s12916-025-04481-5"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"226","DOI":"10.1111\/nbu.12278","article-title":"The role of polyphenols in modern nutrition","volume":"42","author":"Williamson","year":"2017","journal-title":"Nutr. Bull."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Grosso, G., Godos, J., Currenti, W., Micek, A., Falzone, L., Libra, M., Giampieri, F., Forbes-Hern\u00e1ndez, T.Y., Quiles, J.L., and Battino, M. (2022). The effect of dietary polyphenols on vascular health and hypertension: Current evidence and mechanisms of action. Nutrients, 14.","DOI":"10.3390\/nu14030545"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Carresi, C., Gliozzi, M., Musolino, V., Scicchitano, M., Scarano, F., Bosco, F., Nucera, S., Maiuolo, J., Macr\u00ec, R., and Ruga, S. (2020). The effect of natural antioxidants in the development of metabolic syndrome: Focus on bergamot polyphenolic fraction. Nutrients, 12.","DOI":"10.3390\/nu12051504"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Vedova, D.L., Gado, F., Vieira, T.A., Grandini, N.A., Pal\u00e1cio, T.L.N., Siqueira, J.S., Carini, M., Bombardelli, E., Correa, C.R., and Aldini, G. (2023). Chemical, nutritional and biological evaluation of a sustainable and scalable complex of phytochemicals from bergamot by-products. Molecules, 28.","DOI":"10.3390\/molecules28072964"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"C\u00eempeanu, I.-A., Boru, C., Dehelean, C.A., Liga, S., Cosoroab\u0103, R.M., Ardelean, S., Popescu, R., and Vlad, D. (2026). Melissa officinalis L. (Lemon Balm): An integrative review of phytochemistry and evidence from preclinical research to clinical studies. Plants, 15.","DOI":"10.3390\/plants15040650"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Mathews, I.M., Eastwood, J., Lamport, D.J., Cozannet, R.L., Fanca-Berthon, P., and Williams, C.M. (2024). Clinical Efficacy and Tolerability of Lemon Balm (Melissa officinalis L.) in Psychological Well-Being: A Review. Nutrients, 16.","DOI":"10.3390\/nu16203545"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Nowak, M., Sasak, K., Wlodarczyk, A., Grabska-Kobylecka, I., Sarniak, A., and Nowak, D. (2025). AModified Fenton\u2019s system Fe2+-EGTA\u2013H2O2 reveals that redox activities of simple polyphenols are suppressed in binary mixtures. Molecules, 30.","DOI":"10.3390\/molecules30112269"},{"key":"ref_40","first-page":"1975","article-title":"Free radical scavenging activity of polyphenols","volume":"7","author":"Yunusa","year":"2015","journal-title":"J. Chem. Pharm. Res."},{"key":"ref_41","unstructured":"Fotyma, M., and Mercik, S. (2003). Agricultural Chemistry, PWRL."},{"key":"ref_42","unstructured":"Dobrza\u0144ski, B., and Zawadzki, S. (1995). Soil Science, PWRL."},{"key":"ref_43","unstructured":"European Union (1986). Council Directive 86\/278\/EEC of 12 June on the protection of the environment, and in particular of the soil, when sewage sludge is used in agriculture. Off. J. L, 181, 6\u201312."},{"key":"ref_44","unstructured":"IUSS Working Group WRB (2006). World Reference Base for Soil Resources World Soil Resources Reports No. 103, FAO."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1007\/s11270-017-3508-0","article-title":"Heavy metal uptake by herbs. V. Metal accumulation and physiological effects induced by thiuram in Ocimum basilicum L.","volume":"228","author":"Lisowska","year":"2017","journal-title":"Water Air Soil Pollut."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Geisler, M., and Venema, K. (2011). Iron transport and signaling in plants. Transporters and Pumps in Plant Signalin, Springer.","DOI":"10.1007\/978-3-642-14369-4"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Seregin, I.V., and Kozhevnikova, A.D. (2025). Plant responses to heavy metals: From deficiency to excess. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26052064"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Hu, G., Zhou, X., Chao, M., and Hu, H. (2024). Roles of exogenous 5-aminolevulinic acid and dihydroporphyrin iron in chlorophyll precursor synthesis and chlorophyll-related enzyme activities in wheat under different light intensities. Agronomy, 14.","DOI":"10.3390\/agronomy14112464"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1104\/pp.69.1.107","article-title":"Effects of iron and oxygen on chlorophyll biosynthesis","volume":"69","author":"Spiller","year":"1982","journal-title":"Plant Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"381","DOI":"10.32615\/ps.2024.038","article-title":"Chloroplast antioxidant reactions associated with zinc-alleviating effects on iron toxicity in wheat seedlings","volume":"62","author":"Xu","year":"2024","journal-title":"Photosynthetica"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Kastanek, F., Spacilova, M., Krystynik, P., Dlaskova, M., and Solcova, O. (2023). Fenton reaction\u2013unique but still mysterious. Processes, 11.","DOI":"10.3390\/pr11020432"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"710093","DOI":"10.3389\/fpls.2021.710093","article-title":"Iron deficiency leads to chlorosis through impacting chlorophyll synthesis and nitrogen metabolism in Areca catechu L.","volume":"12","author":"Li","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2216","DOI":"10.1111\/pce.12339","article-title":"Membrane transporters mediating root signaling and adaptive responses to oxygen deprivation and soil flooding","volume":"37","author":"Shabala","year":"2014","journal-title":"Plant Cell Environ."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Wang, Y., Kang, Y., Zhong, M., Zhang, L., Chai, X., Jiang, X., and Yang, X. (2022). Effects of iron deficiency stress on plant growth and quality in flowering Chinese cabbage and its adaptive response. Agronomy, 12.","DOI":"10.3390\/agronomy12040875"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1243","DOI":"10.1007\/s10311-020-01012-x","article-title":"Root exudates ameliorate cadmium tolerance in plants: A review","volume":"18","author":"Bali","year":"2020","journal-title":"Environ. Chem. Lett."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/j.envpol.2016.05.080","article-title":"Effect of heavy metals and organic matter on root exudates (low molecular weight organic acids) of herbaceous species: An assessment in sand and soil conditions under different levels of contamination","volume":"216","year":"2016","journal-title":"Environ. Pollut."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1023\/A:1004865811529","article-title":"Cadmium mobilization and plant availability\u2013the impact of organic acids commonly exuded from roots","volume":"230","author":"Nigam","year":"2001","journal-title":"Plant Soil"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.plantsci.2006.03.005","article-title":"Phenolics metabolism and lignin synthesis in root suspension cultures of Panax ginseng in response to copper stress","volume":"171","author":"Ali","year":"2006","journal-title":"Plant Sci."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"100038","DOI":"10.1016\/j.stress.2021.100038","article-title":"Functional aspects of plant secondary metabolites in metal stress tolerance and their importance in pharmacology","volume":"2","author":"Anjitha","year":"2021","journal-title":"Plant Stress"},{"key":"ref_60","unstructured":"(2007). Soil Quality\u2014Sampling\u2014Part 4: Rules for Procedure During the Research Areas of Natural, Semi-Natural and Cultivated (Standard No. PN-ISO 10381-4)."},{"key":"ref_61","unstructured":"(1997). Agricultural Chemical Analysis of the Soil. Determination of pH (Standard No. PN-ISO 10390)."},{"key":"ref_62","unstructured":"(2000). Standard Test Methods for Moisture, Ash, and Organic Matter of Peat and Other Organic Soils (Standard No. ASTMD2974-00)."},{"key":"ref_63","unstructured":"Schumacher, B.A. (2002). Methods for the Determination of Total Organic Carbon (TOC) in Soils and Sediments, United States Environmental Protection Agency, Environmental Sciences Division, National Exposure Research Laboratory."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1023\/B:JRNC.0000020909.67144.fc","article-title":"Preparation and preliminary certification of two new Polish CRMs for inorganic trace analysis","volume":"259","author":"Danko","year":"2004","journal-title":"J. Radioanal. Nucl. Chem."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/j.foodchem.2006.06.066","article-title":"Phenolic composition of Flacourtia indica, Opuntia megacantha and Sclerocarya birrea","volume":"103","author":"Ndhlala","year":"2007","journal-title":"Food Chem."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Goodson, D.Z. (2011). Mathematical Methods for Physical and Analytical Chemistry, Wiley.","DOI":"10.1002\/9781118135204"},{"key":"ref_67","first-page":"21","article-title":"Power comparisons of Shapiro-Wilk, Kolmogorov-Smirnov, Lilliefors and Anderson-Darling tests","volume":"2","author":"Razali","year":"2011","journal-title":"J. Stat. Model. Anal."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/14\/6211\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T04:27:05Z","timestamp":1784176025000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/14\/6211"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,11]]},"references-count":67,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2026,7]]}},"alternative-id":["ijms27146211"],"URL":"https:\/\/doi.org\/10.3390\/ijms27146211","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,11]]}}}