{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T17:08:54Z","timestamp":1785431334683,"version":"3.56.0"},"reference-count":82,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2026,6,17]],"date-time":"2026-06-17T00:00:00Z","timestamp":1781654400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"University of Tabriz, International and Academic Cooperation Directorate","award":["1450\/2021-01-20"],"award-info":[{"award-number":["1450\/2021-01-20"]}]},{"award":["1450\/2021-01-20"],"award-info":[{"award-number":["1450\/2021-01-20"]}],"id":[{"id":"https:\/\/ror.org\/01papkj44","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004837","name":"Spanish MICIN","doi-asserted-by":"publisher","award":["PID2019-104000RB-I00"],"award-info":[{"award-number":["PID2019-104000RB-I00"]}],"id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Serbian Ministry of Science, Technological Development and Innovations","award":["451-03-33\/2026-03\/200053"],"award-info":[{"award-number":["451-03-33\/2026-03\/200053"]}]},{"award":["451-03-33\/2026-03\/200053"],"award-info":[{"award-number":["451-03-33\/2026-03\/200053"]}],"id":[{"id":"https:\/\/ror.org\/01znas443","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Bryophytes are key components of acid\u2013soil ecosystems; however, their capacity for aluminum (Al) accumulation and tolerance remains poorly understood. In this study, bryophytes and a limited number of pteridophyte and lichen species were collected from acidic soils of tea plantations and adjacent forest stands in the Caspian region of northern Iran and analyzed. Nearly all bryophyte specimens exhibited Al concentrations above the critical accumulation threshold (1000 \u00b5g g\u22121 DW), with some reaching values exceeding 28,000 \u00b5g g\u22121 DW, confirming their strong accumulation capacity. After Al, iron was the most abundantly accumulated metal (1430\u201322,800 \u00b5g g\u22121 DW), followed by manganese (100\u20133100 \u00b5g g\u22121 DW). The sampled lichen species accumulated Al at concentrations between 1063 and 9154 \u00b5g g\u22121 DW, while Al levels in the aerial parts of pteridophytes rarely exceeded the critical threshold; when they did, accumulation occurred predominantly in old and fertile fronds rather than sterile ones. Three field-collected bryophyte species\u2014Barbula unguiculata, Palamocladium euchloron, and Hypnum cupressiforme\u2014were acclimated to laboratory conditions and treated with two Al levels (without or with 150 \u00b5M Al, pH 4.0) for 12 weeks. The leafy shoots were analyzed for their antioxidant response, osmolyte accumulation, phenolic metabolism, callose deposition, and carboxylic-acid profile. Histochemical analyses revealed predominant localization of Al in cell walls, associated with enrichment of pectin and uronic acids. These responses were most pronounced in H. cupressiforme, followed by P. euchloron, and least evident in B. unguiculata. Elevated levels of intracellular detoxification compounds\u2014phenolics, flavonoids, and carboxylic acids (tartaric, oxalic, malic, and citric acids)\u2014were detected, again with species-specific differences. Overall, the results reveal that bryophytes employ multiple physiological strategies to tolerate Al toxicity, with substantial interspecific variation. These findings emphasize their ecological significance and provide a foundation for future research on the physiological and evolutionary mechanisms underlying Al tolerance and accumulation in early land plants.<\/jats:p>","DOI":"10.3390\/plants15121877","type":"journal-article","created":{"date-parts":[[2026,6,17]],"date-time":"2026-06-17T08:22:56Z","timestamp":1781684576000},"page":"1877","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Bryophytes as Strong Aluminum Accumulators in Acidic Soils: Cell-Wall Binding and Physiological Tolerance Mechanisms"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0552-4156","authenticated-orcid":false,"given":"Roghieh","family":"Hajiboland","sequence":"first","affiliation":[{"name":"Department of Plant, Cell and Molecular Biology, University of Tabriz, Tabriz 51666-16471, Iran"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6730-475X","authenticated-orcid":false,"given":"Aiuob","family":"Moradi","sequence":"additional","affiliation":[{"name":"Forests and Rangelands Research Department, Guilan Agricultural and Education Center, Agricultural Research, Education and Extension Organization (AREEO), Rasht 41635-3394, Iran"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hedieh","family":"Majmoueh-Koub","sequence":"additional","affiliation":[{"name":"Department of Plant, Cell and Molecular Biology, University of Tabriz, Tabriz 51666-16471, Iran"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8513-3520","authenticated-orcid":false,"given":"Roser","family":"Tolr\u00e0","sequence":"additional","affiliation":[{"name":"Plant Physiology Laboratory, Bioscience Faculty, Universidad Aut\u00f3noma de Barcelona, 08193 Bellaterra, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ana","family":"Paravinja","sequence":"additional","affiliation":[{"name":"Institute for Multidisciplinary Research, National Institute of the Republic of Serbia, University of Belgrade, 11030 Belgrade, Serbia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Milos","family":"Stanojevic","sequence":"additional","affiliation":[{"name":"Institute for Multidisciplinary Research, National Institute of the Republic of Serbia, University of Belgrade, 11030 Belgrade, Serbia"},{"name":"Faculty of Sciences and Mathematics, University of Pristina, 38220 Kosovska Mitrovica, Serbia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9643-138X","authenticated-orcid":false,"given":"Miroslav","family":"Nikolic","sequence":"additional","affiliation":[{"name":"Institute for Multidisciplinary Research, National Institute of the Republic of Serbia, University of Belgrade, 11030 Belgrade, Serbia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3818-0874","authenticated-orcid":false,"given":"Charlotte","family":"Poschenrieder","sequence":"additional","affiliation":[{"name":"Plant Physiology Laboratory, Bioscience Faculty, Universidad Aut\u00f3noma de Barcelona, 08193 Bellaterra, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,6,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1146\/annurev-arplant-043014-114822","article-title":"Plant adaptation to acid soils: The molecular basis for crop aluminum resistance","volume":"66","author":"Kochian","year":"2015","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"N\u00fa\u00f1ez-Delgado, A. (2024). Acidic soils. Planet Earth: Scientific Proposals to Solve Urgent Issues, Springer.","DOI":"10.1007\/978-3-031-53208-5"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1016\/j.scitotenv.2008.06.003","article-title":"A glance into aluminum toxicity and resistance in plants","volume":"400","author":"Poschenrieder","year":"2008","journal-title":"Sci. Total Environ."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1093\/aob\/mcg071","article-title":"A comparative study of metal levels in leaves of some Al-accumulating Rubiaceae","volume":"91","author":"Jansen","year":"2003","journal-title":"Ann. Bot."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1663\/0006-8101(2002)068[0235:AHIAAR]2.0.CO;2","article-title":"Aluminum hyperaccumulation in angiosperms: A review of its phylogenetic significance","volume":"68","author":"Jansen","year":"2002","journal-title":"Bot. Rev."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1093\/aob\/mcf142","article-title":"Aluminium accumulation in leaves of 127 species in Melastomataceae, with comments on the order Myrtales","volume":"90","author":"Jansen","year":"2002","journal-title":"Ann. Bot."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Hajiboland, R., Moradi, A., Kahneh, E., Poschenrieder, C., Nazari, F., Pavlovic, J., Tolr\u00e0, R., Salehi-Lisar, S.Y., and Nikolic, M. (2023). Weed species from tea gardens as a source of novel aluminum hyperaccumulators. Plants, 12.","DOI":"10.3390\/plants12112129"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Panda, S., and Balu\u0161ka, F. (2015). Mechanisms of hyper-resistance and hyper-tolerance to aluminum in plants. Aluminum Stress Adaptation in Plants, Springer.","DOI":"10.1007\/978-3-319-19968-9"},{"key":"ref_9","unstructured":"Gupta, R.K., and Kumar, M. (2013). Bryophytes and ecosystem. Biodiversity of Lower Plants, IK International Publishing House."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"2716","DOI":"10.1890\/09-2095.1","article-title":"Similar cation exchange capacities among bryophyte species refute a presumed mechanism of peatland acidification","volume":"91","author":"Soudzilovskaia","year":"2010","journal-title":"Ecology"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.envpol.2012.03.018","article-title":"Bioacumulation and ultrastructural effects of Cd, Cu, Pb and Zn in the moss Scorpiurum circinatum (Brid.) Fleisch. & Loeske","volume":"166","author":"Basile","year":"2012","journal-title":"Environ. Pollut."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/S0269-7491(00)00227-X","article-title":"Response of mosses to the heavy metal deposition in Poland\u2014An overview","volume":"114","year":"2001","journal-title":"Environ. Pollut."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Murthy, H.N. (2023). Bryophytes as an accumulator of toxic elements from the environment: Recent advances. Bioactive Compounds in Bryophytes and Pteridophytes, Springer.","DOI":"10.1007\/978-3-031-23243-5"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1006\/eesa.1999.1798","article-title":"Uptake of heavy metals to the extracellular and intracellular compartments in three species of aquatic bryophyte","volume":"44","author":"Carballeira","year":"1999","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"3967","DOI":"10.1007\/s00344-022-10866-0","article-title":"Aluminum toxicity in plants: Present and future","volume":"42","author":"Hajiboland","year":"2023","journal-title":"J. Plant Growth Regul."},{"key":"ref_16","first-page":"315","article-title":"An annotated and updated checklist of Iranian bryoflora","volume":"25","author":"Akhani","year":"2004","journal-title":"Cryptogam. Bryol."},{"key":"ref_17","first-page":"231","article-title":"Plant biodiversity of Hyrcanian relict forests, N Iran: An overview of the flora, vegetation, palaeoecology and conservation","volume":"42","author":"Akhani","year":"2010","journal-title":"Pak. J. Bot."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.pld.2021.07.005","article-title":"Plant diversity of Hyrcanian relict forests: An annotated checklist, chorology and threat categories of endemic and near endemic vascular plant species","volume":"44","author":"Ghorbanalizadeh","year":"2022","journal-title":"Plant Divers."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"200085","DOI":"10.1016\/j.teadva.2023.200085","article-title":"Lichens as effective bioindicators for monitoring environmental changes: A comprehensive review","volume":"9","author":"Thakur","year":"2024","journal-title":"Total Environ. Adv."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.geoderma.2005.07.002","article-title":"Competitive adsorption and desorption of copper and zinc in acid soils","volume":"133","author":"Arias","year":"2006","journal-title":"Geoderma"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1016\/j.geoderma.2009.06.005","article-title":"Calcium and magnesium releasing capacity of Alfisols under tea in south India","volume":"152","author":"Senthurpandian","year":"2009","journal-title":"Geoderma"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"11412","DOI":"10.1038\/s41598-022-14851-1","article-title":"Acidic condition accelerates cation release from purple rock in Southwestern China","volume":"12","author":"Zhao","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1146\/annurev.arplant.55.031903.141655","article-title":"How do crop plants tolerate acid soils? Mechanisms of aluminum tolerance and phosphorous efficiency","volume":"55","author":"Kochian","year":"2004","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_24","unstructured":"Barker, A.V., and Pilbeam, D.J. (2007). Molybdenum. Handbook of Plant Nutrition, Taylor Francis Group."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1765","DOI":"10.1111\/ppl.13382","article-title":"Alleviating aluminum toxicity in plants: Implications of reactive oxygen species signaling and crosstalk with other signaling pathways","volume":"173","author":"Ranjan","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"4386","DOI":"10.1007\/s12033-024-01351-y","article-title":"Intertwining of cellular osmotic stress handling mechanisms and heavy metal accumulation","volume":"67","author":"Saavedra","year":"2025","journal-title":"Mol. Biotechnol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1093\/jxb\/erj027","article-title":"Involvement of soluble sugars in reactive oxygen species balance and responses to oxidative stress in plants","volume":"57","author":"Sulmon","year":"2006","journal-title":"J. Exp. Bot."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1111\/plb.13363","article-title":"Proline, a multifaceted signalling molecule in plant responses to abiotic stress: Understanding the physiological mechanisms","volume":"24","author":"Ghosh","year":"2022","journal-title":"Plant Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1007\/s10265-016-0836-x","article-title":"Dehydration-responsive features of Atrichum undulatum","volume":"129","author":"Hu","year":"2016","journal-title":"J. Plant Res."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e12419","DOI":"10.7717\/peerj.12419","article-title":"Acclimation of liverwort Marchantia polymorpha to physiological drought reveals important roles of antioxidant enzymes, proline and abscisic acid in land plant adaptation to osmotic stress","volume":"9","author":"Ghosh","year":"2021","journal-title":"Peer J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"16401","DOI":"10.1038\/s41598-020-73358-9","article-title":"Al exposure increases proline levels by different pathways in an Al-sensitive and an Al-tolerant rye genotype","volume":"10","author":"AbdElgawad","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Lin, D., Li, Q., Suleiman, M., An, M., and Zhu, Y. (2026). Microbiome and metabolomics analysis of different bryophytes under cadmium stress. BMC Plant Biol., 26.","DOI":"10.1186\/s12870-026-08651-2"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/S1360-1385(01)01961-6","article-title":"Aluminium tolerance in plants and the complexing role of organic acids","volume":"6","author":"Ma","year":"2001","journal-title":"Trends Plant Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"3025","DOI":"10.1021\/acs.jafc.6b00331","article-title":"Polyphenol\u2013aluminum complex formation: Implications for aluminum tolerance in plants","volume":"64","author":"Zhang","year":"2016","journal-title":"J. Agric. Food Chem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"4396","DOI":"10.1093\/jxb\/erac085","article-title":"Stress, senescence, and specialized metabolites in bryophytes","volume":"73","author":"Kulshrestha","year":"2022","journal-title":"J. Exp. Bot."},{"key":"ref_36","first-page":"523","article-title":"Phenolic compounds and their antioxidant activity in plants growing under heavy metal stress","volume":"15","author":"Michalak","year":"2006","journal-title":"Pol. J. Environ. Stud."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"709","DOI":"10.4161\/psb.6.5.15069","article-title":"Stress-induced flavonoid biosynthesis and the antioxidant machinery of plants","volume":"6","author":"Fini","year":"2011","journal-title":"Plant Signal. Behav."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Burbidge, C.A., Ford, C.M., Melino, V.J., Wong, D.C.J., Jia, Y., Jenkins, C.L.D., Soole, K.L., Castellarin, S.D., Darriet, P., and Rienth, M. (2021). Biosynthesis and cellular functions of tartaric acid in grapevines. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.643024"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1631\/jzus.B1900188","article-title":"Mechanisms and regulation of aluminum-induced secretion of organic acid anions from plant roots","volume":"20","author":"Yang","year":"2019","journal-title":"J. Zhejiang Univ. Sci. B"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2531","DOI":"10.1080\/10643389.2013.829764","article-title":"Enhanced phytoremediation: A review of low molecular weight organic acids and surfactants used as amendments","volume":"44","author":"Agnello","year":"2014","journal-title":"Crit. Rev. Environ. Sci. Technol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Pannequin, A., Muselli, A., Marcourt, L., Ferreira-Queiroz, E., Quiros-Guerrero, L.M., Asakawa, Y., Dounoue-Kubo, M., and Wolfender, J.L. (2025). Comprehensive comparative metabolome study of a large collection of Corsican bryophytes. Front. Plant Sci., 15.","DOI":"10.3389\/fpls.2024.1470307"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Zhu, Y., Lin, D., Li, Q., An, M., and Lv, J. (2025). Metabolomic analysis of the responses of bryophyte Tortella tortuosa (Hedw.) Limpr. to cadmium (Cd) stress. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26072856"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Murthy, H.N. (2023). Bioactive compounds of pteridophytes. Bioactive Compounds in Bryophytes and Pteridophytes, Springer.","DOI":"10.1007\/978-3-031-23243-5"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1111\/nph.13011","article-title":"Physiological characterization of aluminum tolerance and accumulation in tartary and wild buckwheat","volume":"205","author":"Wang","year":"2015","journal-title":"New Phytol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1104\/pp.113.4.1033","article-title":"Internal detoxification mechanism of Al in hydrangea (identification of Al form in the leaves)","volume":"113","author":"Ma","year":"1997","journal-title":"Plant Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Horst, W.J., Schenk, M.K., B\u00fcrkert, A., Claassen, N., Flessa, H., Frommer, W.B., Goldbach, H., Olfs, H.W., R\u00f6mheld, V., and Sattelmacher, B. (2001). Effect of aluminium on exudation of organic acid anions in tea plants. Plant Nutrition: Food Security and Sustainability of Agro-Ecosystems Through Basic and Applied Research, Springer.","DOI":"10.1007\/0-306-47624-X"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Dumont, S., and Rivoal, J. (2019). Consequences of oxidative stress on plant glycolytic and respiratory metabolism. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.00166"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.wem.2011.01.001","article-title":"Mechanism of action of stinging nettles","volume":"22","author":"Cummings","year":"2011","journal-title":"Wilderness Environ. Med."},{"key":"ref_49","unstructured":"Laurila, T., and Lindfors, V. (1999). Exchange of oxygenated volatile organic compounds between boreal lichens and the atmosphere. Biogenic VOC Emissions and Photochemistry in the Boreal Regions of Europe, CEC Air Pollution Research Report, Official Publication of the European Commission."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"3041","DOI":"10.1093\/jxb\/erw090","article-title":"Photorespiratory glycolate\u2013glyoxylate metabolism","volume":"67","author":"Dellero","year":"2016","journal-title":"J. Exp. Bot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1007\/s11738-012-1103-y","article-title":"Callose: The plant cell wall polysaccharide with multiple biological functions","volume":"35","year":"2013","journal-title":"Acta Physiol. Plant."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"991","DOI":"10.1104\/pp.124.3.991","article-title":"Aluminum-induced 1\u21923-\u03b2-d-glucan inhibits cell-to-cell trafficking of molecules through plasmodesmata. A new mechanism of aluminum toxicity in plants","volume":"124","author":"Sivaguru","year":"2000","journal-title":"Plant Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s11738-010-0581-z","article-title":"The cell wall in plant cell response to trace metals: Polysaccharide remodeling and its role in defense strategy","volume":"33","year":"2011","journal-title":"Acta Physiol. Plant."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1080\/17429145.2012.712723","article-title":"Increasing in ROS levels and callose deposition in peduncle vascular bundles of wheat (Triticum aestivum L.) grown under nitrogen deficiency","volume":"8","author":"Kong","year":"2013","journal-title":"J. Plant Interact."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1442","DOI":"10.1007\/s00344-025-11955-6","article-title":"Manganese toxicity tolerance in Lamium album and Mentha aquatica, two Lamiaceae species adapted to acidic soils","volume":"45","author":"Nazari","year":"2026","journal-title":"J. Plant Growth Regul."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1213","DOI":"10.1093\/jxb\/eri115","article-title":"Root cell patterning: A primary target for aluminium toxicity in maize","volume":"56","author":"Doncheva","year":"2005","journal-title":"J. Exp. Bot."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1080\/00380768.2018.1553484","article-title":"Aluminum toxicity in plant cells: Mechanisms of cell death and inhibition of cell elongation","volume":"665","author":"Yamamoto","year":"2019","journal-title":"Soil Sci. Plant Nutr."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1093\/aob\/mcq254","article-title":"Genotypic differences in Al resistance and the role of cell-wall pectin in Al exclusion from the root apex in Fagopyrum tataricum","volume":"107","author":"Yang","year":"2011","journal-title":"Ann. Bot."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/j.envexpbot.2008.05.006","article-title":"Pectinous cell wall thickenings formation\u2014A response of moss protonemata cells to lead","volume":"65","author":"Lenartowska","year":"2009","journal-title":"Environ. Exp. Bot."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Petschinger, K., Adlassnig, W., Sabovljevic, M.S., and Lang, I. (2021). Lamina cell shape and cell wall thickness are useful indicators for metal tolerance\u2014An example in bryophytes. Plants, 10.","DOI":"10.3390\/plants10020274"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Boj\u00f3rquez-Quintal, E., Escalante-Maga\u00f1a, C., Echevarr\u00eda-Machado, I., and Mart\u00ednez-Est\u00e9vez, M. (2017). Aluminum, a friend or foe of higher plants in acid soils. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01767"},{"key":"ref_62","first-page":"91","article-title":"Application of the global bioclimatic classification to Iran: Implications for understanding the modern vegetation and biogeography","volume":"37","author":"Djamali","year":"2011","journal-title":"Ecol. Mediterr."},{"key":"ref_63","unstructured":"Rechinger, K.H. (1963\u20132010). Flora Iranica, Naturhistorisches Museum Wien."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Smith, A.J.E. (2004). The Moss Flora of Britain and Ireland, Cambridge University Press.","DOI":"10.1017\/CBO9780511541858"},{"key":"ref_65","unstructured":"K\u00fcrschner, H., and Frey, W. (2011). Liverworts, Mosses and Hornworts of Southwest Asia (Marchantiophyta, Bryophyta, Anthocerotophyta), J. Cramer. Nova Hedwigia (Supplement 139)."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"188","DOI":"10.2134\/jeq2007.0034","article-title":"Titanium as an indicator of residual soil on arid-land plants","volume":"38","author":"Cook","year":"2009","journal-title":"J. Environ. Qual."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"2329","DOI":"10.1007\/s10530-024-03318-1","article-title":"Effect of aluminum on growth and herbicide resistance in Commelina communis and Tradescantia fluminensis, two invasive weed species in tea gardens","volume":"26","author":"Hajiboland","year":"2024","journal-title":"Biol. Invasions"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.envexpbot.2013.01.001","article-title":"Differential aluminum resistance in Brachiaria species","volume":"89","author":"Arroyave","year":"2013","journal-title":"Environ. Exp. Bot."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"86","DOI":"10.32604\/phyton.2015.84.086","article-title":"Localization and compartmentation of Al in the leaves and roots of tea plants","volume":"84","author":"Hajiboland","year":"2015","journal-title":"Phyton"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Sheng, H., Zeng, J., Liu, Y., Wang, X., Wang, Y., Kang, H., Fan, X., Sha, L., Zhang, H., and Zhou, Y. (2016). Sulfur mediated alleviation of Mn toxicity in polish wheat relates to regulating Mn allocation and improving antioxidant system. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01382"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1042\/bst0110591","article-title":"Determination of total carotenoids and chlorophylls a and b of leaf in different solvents","volume":"11","author":"Lichtenthaler","year":"1983","journal-title":"Biochem. Soc. Trans."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1007\/978-1-60761-702-0_22","article-title":"Extraction and analysis of soluble carbohydrates","volume":"Volume 639","author":"Sunkar","year":"2010","journal-title":"Plant Stress Tolerance. Methods in Molecular Biology"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/BF00018060","article-title":"Rapid determination of free proline for water-stress studies","volume":"39","author":"Bates","year":"1973","journal-title":"Plant Soil"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"730","DOI":"10.4236\/ajac.2014.511081","article-title":"Optimized assay for hydrogen peroxide determination in plant tissue using potassium iodide","volume":"5","author":"Junglee","year":"2014","journal-title":"Am. J. Anal. Chem."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1007\/s004250050524","article-title":"Improving the thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds","volume":"207","author":"Hodges","year":"1999","journal-title":"Planta"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1452","DOI":"10.1104\/pp.17.01608","article-title":"Biochemical and structural analysis of substrate specificity of a phenylalanine ammonia-lyase","volume":"176","author":"Jun","year":"2018","journal-title":"Plant Physiol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.foodchem.2003.10.017","article-title":"Partial purification and characterization of polyphenol oxidase from artichoke (Cynara scolymus L.) heads","volume":"87","author":"Aydemir","year":"2004","journal-title":"Food Chem."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"6852","DOI":"10.3390\/molecules18066852","article-title":"Application and analysis of the folin ciocalteu method for the determination of the total phenolic content from Limonium brasiliense L.","volume":"18","author":"Blainski","year":"2013","journal-title":"Molecules"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1016\/j.ijbiomac.2013.05.020","article-title":"Antioxidant and antimicrobial activities of Lycium shawii fruits extract","volume":"60","author":"Dahech","year":"2013","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1007\/s11104-020-04599-w","article-title":"Selenium activates components of iron acquisition machinery in oilseed rape roots","volume":"452","author":"Hajiboland","year":"2020","journal-title":"Plant Soil"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1940","DOI":"10.1104\/pp.100.4.1940","article-title":"Fourier transform infrared microspectroscopy is a new way to look at plant cell walls","volume":"100","author":"Hammouri","year":"1992","journal-title":"Plant Physiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1046\/j.1365-313X.2003.01807.x","article-title":"Classification and identification of Arabidopsis cell wall mutants using Fourier-Transform InfraRed (FT-IR) microspectroscopy","volume":"35","author":"Mouille","year":"2003","journal-title":"Plant J."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/12\/1877\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,17]],"date-time":"2026-06-17T09:42:55Z","timestamp":1781689375000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/12\/1877"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,17]]},"references-count":82,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2026,6]]}},"alternative-id":["plants15121877"],"URL":"https:\/\/doi.org\/10.3390\/plants15121877","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,17]]}}}