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The mechanisms of humic acid (HA) to alleviate Al3+ stress in plants remain unclear. \u2018Sour pummelo\u2019 (Citrus grandis (L.) Osbeck) seedlings were exposed to 0.5 (HA0.5), 0.1 (HA0.1), or 0 (HA0) mM sodium humate and 1.2 (Al1.2) or 0 (Al0) mM AlCl3\u00b76H2O for 128 days. Thereafter, the research examined biomass; Al and mineral nutrients; leaf photosynthetic performance; and leaf and root nonstructural carbohydrates, reactive oxygen species metabolism, and related physiological parameters. Al1.2 significantly reduced whole plant dry weight (DW), root DW, leaf CO2 assimilation (ACO2), and chlorophyll a + b concentration by 61%, 45%, 61%, and 35%, respectively, at HA0, but only 48%, 17%, 44%, and 11%, respectively, at HA0.5. Further analysis suggested that the addition of HA endowed Citrus with Al resilience by the following several aspects: (a) lessened tissue (leaf, stem, and root) concentrations of Al and enhanced capacity to maintain macronutrient (S, K, Mg, Ca, N, and P) homeostasis at Al1.2; (b) improved capacity to combat oxidative stress at Al1.2; and (c) enhanced ACO2 and growth at Al1.2. Further analysis indicated that HA-mediated alleviation of growth decline caused by Al1.2 involved (a) reduced ability to absorb Al and less root-to-shoot Al transport and (b) increased ability to maintain macronutrient homeostasis and to combat oxidative stress; and that HA-mediated alleviation of leaf chlorophyll and ACO2 decline and photosynthetic electron transport chain impairment involved less leaf Al concentration and improved leaf macronutrient homeostasis. To conclude, the addition of HA lowered roots\u2019 ability to absorb Al and tissue Al concentration and subsequently mitigated Al-toxic impairment to root growth and function, thereby enhancing the ability of plants to maintain macronutrient homeostasis, and hence alleviating Al1.2-stimulated oxidative damage and inhibition of ACO2 and growth.<\/jats:p>","DOI":"10.3390\/plants15152370","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T06:17:40Z","timestamp":1785737860000},"page":"2370","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Humic Acid Alleviates Aluminum Toxicity in Citrus grandis (L.) Osbeck: Insight from Growth, Gas Exchange, and Related Physiological Parameters"],"prefix":"10.3390","volume":"15","author":[{"given":"Qian","family":"Shen","sequence":"first","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tian-Tian","family":"Xia","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liang-Yuan","family":"Tong","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bin-Bin","family":"Lan","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei-Lin","family":"Huang","sequence":"additional","affiliation":[{"name":"College of Environmental and Biological Engineering, Putian University, Putian 351100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ti","family":"Wu","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"College of Environmental and Biological Engineering, Putian University, Putian 351100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xin","family":"Ye","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ning-Wei","family":"Lai","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8425-1306","authenticated-orcid":false,"given":"Li-Song","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Resources and Environment, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,31]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"101200","DOI":"10.1016\/j.xplc.2024.101200","article-title":"Aluminum resistance in plants: A critical review focusing on STOP1","volume":"6","author":"Huang","year":"2025","journal-title":"Plant Commun."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1146\/annurev.pp.46.060195.001321","article-title":"Cellular mechanisms of aluminum toxicity and resistance in plants","volume":"46","author":"Kochian","year":"1995","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Wang, F., Li, X., Chen, C., Zhao, L., and Wei, Y. (2025). Mechanism of melatonin in alleviating aluminum toxicity in plants: A review. Biology, 14.","DOI":"10.3390\/biology14101316"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1008","DOI":"10.1126\/science.1182570","article-title":"Significant acidification in major Chinese croplands","volume":"327","author":"Guo","year":"2010","journal-title":"Science"},{"key":"ref_5","first-page":"9","article-title":"Ecological region division of soil based supplementary fertilization and decrement fertilization in China Citrus orchards","volume":"41","author":"Wu","year":"2022","journal-title":"J. Huazhong Agri. Univ."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"109366","DOI":"10.1016\/j.plaphy.2024.109366","article-title":"Glutathione mitigates aluminum toxicity in root-apex transition zone of rice through reducing aluminum absorption and maintaining redox balance","volume":"219","author":"Jiang","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Yang, H., Xie, R.R., Xia, T.T., Tong, L.Y., Wu, T., Ye, X., Huang, Z.R., Yang, L.T., and Chen, L.S. (2026). Coumarin reduces aluminum-induced inhibition of growth and photosynthesis in Citrus grandis by reducing tissue Al concentration and maintaining nutrient and redox homeostasis. Plants, 15.","DOI":"10.3390\/plants15111694"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1016\/S0176-1617(00)80106-6","article-title":"Gas exchange and chlorophyll fluorescence in four Citrus rootstocks under aluminum stress","volume":"157","author":"Pereira","year":"2000","journal-title":"J. Plant Physiol."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Yang, M., Tan, L., Xu, Y., Zhao, Y., Cheng, F., Ye, S., and Jiang, W. (2015). Effect of low pH and aluminum toxicity on the photosynthetic characteristics of different fast-growing Eucalyptus vegetatively propagated clones. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0130963"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"572","DOI":"10.2135\/cropsci1986.0011183X002600030030x","article-title":"Photosynthesis, chlorophyll, and transpiration responses in aluminum stressed wheat and sorghum","volume":"26","author":"Ohki","year":"1986","journal-title":"Crop Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1007\/s13205-024-04192-3","article-title":"Evaluating the potential of Acinetobacter calcoaceticus in alleviation of aluminium stress in Triticum aestivum","volume":"15","author":"Okla","year":"2025","journal-title":"3 Biotech."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1080\/01904169409364729","article-title":"Aluminum toxicity in tomato. Part 2. Leaf gas exchange, chlorophyll content, and invertase activity","volume":"17","author":"Simon","year":"1994","journal-title":"J. Plant Nutr."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"117001","DOI":"10.1016\/j.ecoenv.2024.117001","article-title":"Physiological and comprehensive transcriptome analysis reveals distinct regulatory mechanisms for aluminum tolerance of Trifolium repens","volume":"284","author":"Nie","year":"2024","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"114421","DOI":"10.1016\/j.ecoenv.2022.114421","article-title":"Magnesium alleviates aluminum-induced growth inhibition by enhancing antioxidant enzyme activity and carbon-nitrogen metabolism in apple seedlings","volume":"249","author":"Lyu","year":"2023","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1081\/PLN-120002962","article-title":"Responses of the photosynthetic apparatus to aluminum stress in two sorghum cultivars","volume":"25","author":"Peixoto","year":"2002","journal-title":"J. Plant Nutr."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"120313","DOI":"10.1016\/j.envpol.2022.120313","article-title":"Subcellular compartmentalization of aluminum reduced its hazardous impact on rye photosynthesis","volume":"315","author":"AbdElgawad","year":"2022","journal-title":"Environ. Pollut."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1548","DOI":"10.1093\/treephys\/tpy035","article-title":"Aluminum effects on photosynthesis, reactive oxygen species and methylglyoxal detoxification in two Citrus species differing in aluminum tolerance","volume":"38","author":"Guo","year":"2018","journal-title":"Tree Physiol."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Moustaka, J., Ouzounidou, G., Sperdouli, I., and Moustakas, M. (2018). Photosystem II is more sensitive than photosystem I to Al3+ induced phytotoxicity. Materials, 11.","DOI":"10.3390\/ma11091772"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"\u017burek, G., Rybka, K., Pogrzeba, M., Krzy\u017cak, J., and Prokopiuk, K. (2014). Chlorophyll a fluorescence in evaluation of the effect of heavy metal soil contamination on perennial grasses. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0091475"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"9058715","DOI":"10.1155\/2019\/9058715","article-title":"Increasing nutrient solution pH alleviated aluminum-induced inhibition of growth and impairment of photosynthetic electron transport chain in Citrus sinensis seedlings","volume":"2019","author":"Yang","year":"2019","journal-title":"BioMed Res. Int."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Guo, P., Li, Q., Qi, Y.P., Yang, L.T., Ye, X., Chen, H.H., and Chen, L.S. (2017). Sulfur-mediated-alleviation of aluminum-toxicity in Citrus grandis seedlings. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18122570"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1016\/j.envexpbot.2009.07.010","article-title":"Aluminum stress induces up-regulation of an efficient antioxidant system in the Al-tolerant maize line but not in the Al-sensitive line","volume":"67","author":"Giannakoula","year":"2010","journal-title":"Environ. Exp. Bot."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1104\/pp.010417","article-title":"Aluminum toxicity is associated with mitochondrial dysfunction and the production of reactive oxygen species in plant cells","volume":"128","author":"Yamamoto","year":"2002","journal-title":"Plant Physiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1111\/tpj.12290","article-title":"Targeted expression of SbMATE in the root distal transition zone is responsible for sorghum aluminum resistance","volume":"76","author":"Sivaguru","year":"2013","journal-title":"Plant J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S1672-6308(12)60042-0","article-title":"Involvement of antioxidative defense system in rice seedlings exposed to aluminum toxicity and phosphorus deficiency","volume":"19","author":"Guo","year":"2012","journal-title":"Rice Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1278","DOI":"10.1046\/j.0016-8025.2001.00783.x","article-title":"Transgenic Brassica napus plants overexpressing aluminum-induced mitochondrial manganese superoxide dismutase cDNA are resistant to aluminum","volume":"24","author":"Basu","year":"2001","journal-title":"Plant Cell Environ."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1007\/s11103-017-0644-2","article-title":"Overexpression of a peroxidase gene (AtPrx64) of Arabidopsis thaliana in tobacco improves plant\u2019s tolerance to aluminum stress","volume":"95","author":"Wu","year":"2017","journal-title":"Plant Mol. Biol."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Sible, C.N., Seebauer, J.R., and Below, F.E. (2021). Plant biostimulants: A categorical review, their implications for row crop production, and relation to soil health indicators. Agronomy, 11.","DOI":"10.3390\/agronomy11071297"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1016\/j.jenvman.2017.05.073","article-title":"Application of drum compost and vermicompost to improve soil health, growth, and yield parameters for tomato and cabbage plants","volume":"200","author":"Goswami","year":"2017","journal-title":"J. Environ. Manag."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"116958","DOI":"10.1016\/j.ecoenv.2024.116958","article-title":"Biochar combined with humic acid improves the soil environment and regulate microbial communities in apple replant soil","volume":"283","author":"Liu","year":"2024","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.scienta.2015.09.013","article-title":"Humic and fulvic acids as biostimulants in horticulture","volume":"196","author":"Canellas","year":"2015","journal-title":"Sci. Horti."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.pedsph.2022.07.008","article-title":"Humic substances: Structure, function and benefits for agroecosystems: A review","volume":"33","author":"Tiwari","year":"2023","journal-title":"Pedosphere"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"115546","DOI":"10.1016\/j.envpol.2020.115546","article-title":"Regulation mechanisms of humic acid on Pb stress in tea plant (Camellia sinensis L.)","volume":"267","author":"Duan","year":"2020","journal-title":"Environ. Pollut."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"144043","DOI":"10.1016\/j.ijbiomac.2025.144043","article-title":"Role of humic acid for climate change adaptation measures to boost up sustainable agriculture and soil health: A potential review","volume":"313","author":"Antu","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"7581","DOI":"10.1007\/s42729-024-02061-9","article-title":"Soil-plant integrative supplementation with humic acid and antioxidants improves growth, fruit quality, and antioxidant capacity of Cd-stressed Solanum melongena","volume":"24","author":"Sayed","year":"2024","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"122114","DOI":"10.1016\/j.indcrop.2025.122114","article-title":"Mechanistic aspect of humic acid and Pseudomonas nitroreducens strain UF50 to alleviate nickel toxicity in soybean","volume":"237","author":"Odongkara","year":"2025","journal-title":"Ind. Crop Prod."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Huang, W.T., Chen, X.F., Huang, W.L., Shen, Q., Lu, F., Lai, N.W., Guo, J., Yang, L.T., Ye, X., and Chen, L.S. (2025). Humic acid enhances antioxidant and glyoxalase systems to combat copper toxicity in Citrus. Agronomy, 15.","DOI":"10.3390\/agronomy15010099"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1553","DOI":"10.1007\/s11104-025-07816-6","article-title":"Beneficial effects of commercially available preparations of humic substances and mycorrhiza on growth and photosynthesis of sorghum and hemp cultivated on a metal(loid)-polluted field","volume":"516","author":"Sitko","year":"2025","journal-title":"Plant Soil"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.ecoenv.2018.02.071","article-title":"The humic acid-induced changes in the water status, chlorophyll fluorescence and antioxidant defense systems of wheat leaves with cadmium stress","volume":"155","author":"Yildiztugay","year":"2018","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1071\/S98018","article-title":"Effects of humic and fulvic acids on the rhizotoxicity of lanthanum and aluminium to corn","volume":"36","author":"Diatloff","year":"1998","journal-title":"Aust. J. Soil Res."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Harper, S., and Menzies, N. (2023). Phytotoxic effects of Al on root growth are confounded in the presence of fulvic and humic acids. Soil Syst., 7.","DOI":"10.3390\/soilsystems7030068"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1097\/00010694-198601000-00004","article-title":"Effect of humic acid on aluminum toxicity in corn plants","volume":"141","author":"Tan","year":"1986","journal-title":"Soil Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1038\/196085a0","article-title":"Resistance to aluminium and manganese toxicities in plants related to variety and cation-exchange capacity","volume":"196","author":"Vose","year":"1962","journal-title":"Nature"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1111\/j.1399-3054.2009.01288.x","article-title":"Phosphorus alleviates aluminum-induced inhibition of growth and photosynthesis in Citrus grandis seedlings","volume":"137","author":"Jiang","year":"2009","journal-title":"Physiol. Plant."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Jiang, H.X., Tang, N., Zheng, J.G., and Chen, L.S. (2009). Antagonistic actions of boron against inhibitory effects of aluminum toxicity on growth, CO2 assimilation, ribulose-1,5-bisphosphate carboxylase\/oxygenase, and photosynthetic electron transport probed by the JIP-test, of Citrus grandis seedlings. BMC Plant Biol., 9.","DOI":"10.1186\/1471-2229-9-102"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"129275","DOI":"10.1016\/j.jhazmat.2022.129275","article-title":"Boron contributes to excessive aluminum tolerance in trifoliate orange (Poncirus trifoliata (L.) Raf.) by inhibiting cell wall deposition and promoting vacuole compartmentation","volume":"437","author":"Yan","year":"2022","journal-title":"J. Hazard. Mater."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"114303","DOI":"10.1016\/j.envpol.2020.114303","article-title":"Interactive effects of pH and aluminum on the secretion of organic acid anions by roots and related metabolic factors in Citrus sinensis roots and leaves","volume":"262","author":"Yang","year":"2020","journal-title":"Environ. Pollut."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"120473","DOI":"10.1016\/j.chemgeo.2021.120473","article-title":"Enhancement of soil microstructural stability and alleviation of aluminium toxicity in acidic latosols via alkaline humic acid fertiliser amendment","volume":"583","author":"Deng","year":"2021","journal-title":"Chem. Geol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1080\/00207233.2025.2457867","article-title":"Non-microbial biostimulants for plant growth and abiotic stress mitigation: A review of recent scientific innovations","volume":"74","author":"Jmaili","year":"2025","journal-title":"Int. J. Environ. Stud."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Muratore, C., Espen, L., and Prinsi, B. (2021). Nitrogen uptake in plants: The plasma membrane root transport systems from a physiological and proteomic perspective. Plants, 10.","DOI":"10.3390\/plants10040681"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1002\/jpln.202300413","article-title":"Humic acid enhances phosphorus transport in soil and uptake by maize","volume":"187","author":"Rosolem","year":"2024","journal-title":"J. Plant Nutr. Soil Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"3920","DOI":"10.1007\/s00344-024-11359-y","article-title":"Effects of humic acid-copper interactions on growth, nutrient absorption, and photosynthetic performance of Citrus sinensis seedlings in sand culture","volume":"43","author":"Huang","year":"2024","journal-title":"J. Plant Growth Regul."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1901","DOI":"10.1081\/PLN-100107602","article-title":"Phosphorus deficiency is responsible for biomass reduction of triticale in nutrient with aluminum","volume":"24","author":"Quartin","year":"2001","journal-title":"J. Plant Nutr."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1016\/j.plaphy.2015.11.019","article-title":"The interaction of salicylic acid and Ca2+ alleviates aluminum toxicity in soybean (Glycine max L.)","volume":"98","author":"Lan","year":"2016","journal-title":"Plant Physiol. Biochem."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"39025","DOI":"10.1038\/s41598-025-25724-8","article-title":"Effect of exogenously applied amino acids on photosynthetic pigments, metabolites and enzymatic antioxidants in Zea mays L. subjected to salt stress","volume":"15","author":"Ahmad","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"319","DOI":"10.2174\/1872208316666220412133749","article-title":"The effects of amino acids, phenols and protein hydrolysates as biostimulants on sustainable crop production and alleviated stress","volume":"16","author":"Shahrajabian","year":"2022","journal-title":"Recent Pat. Biotechnol."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Huang, W.T., Zheng, Z.C., Hua, D., Chen, X.F., Zhang, J., Chen, H.H., Ye, X., Guo, J.X., Yang, L.T., and Chen, L.S. (2022). Adaptive responses of carbon and nitrogen metabolisms to nitrogen-deficiency in Citrus sinensis seedlings. BMC Plant Biol., 22.","DOI":"10.1186\/s12870-022-03759-7"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"136523","DOI":"10.1016\/j.chemosphere.2022.136523","article-title":"Effects of exogenous taurine on growth, photosynthesis, oxidative stress, antioxidant enzymes and nutrient accumulation by Trifolium alexandrinum plants under manganese stress","volume":"308","author":"Hafeez","year":"2022","journal-title":"Chemosphere"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.plaphy.2020.10.023","article-title":"Copper toxicity affects phosphorus uptake mechanisms at molecular and physiological levels in Cucumis sativus plants","volume":"157","author":"Feil","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Huang, W.T., Xie, Y.Z., Chen, X.F., Zhang, J., Chen, H.H., Ye, X., Guo, J., Yang, L.T., and Chen, L.S. (2021). Growth, mineral nutrients, photosynthesis and related physiological parameters of Citrus in response to nitrogen deficiency. Agronomy, 11.","DOI":"10.3390\/agronomy11091859"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1111\/pce.13935","article-title":"Phosphate or nitrate imbalance induces stronger molecular responses than combined nutrient deprivation in roots and leaves of chickpea plants","volume":"44","author":"Inoue","year":"2021","journal-title":"Plant Cell Environ."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1081\/PLN-100104970","article-title":"The influence of nitrogen source and aluminum on growth and elemental composition of \u2018Nemaguard\u2019 peach seedlings","volume":"24","author":"Graham","year":"2001","journal-title":"J. Plant Nutr."},{"key":"ref_63","first-page":"27","article-title":"Influence of different levels of aluminum on the development of Citrus rootstock \u2018Swingle\u2019 citrumelo (Citrus paradisi Mcf. \u00d7 Poncirus trifoliata Raf.) in nutrient solution","volume":"43","author":"Filho","year":"2000","journal-title":"Braz. Arch. Biol. Techn."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"T\u00f3th, B., Moloi, M.J., Sz\u0151ke, L., Danter, M., and Grusak, M.A. (2021). Cultivar differences in the biochemical and physiological responses of common beans to aluminum stress. Plants, 10.","DOI":"10.3390\/plants10102097"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Lopez, G., Ahmadi, S.H., Amelung, W., Athmann, M., Ewert, F., Gaiser, T., Gocke, M.I., Kautz, T., Postma, J., and Rachmilevitch, S. (2023). Nutrient deficiency effects on root architecture and root-to-shoot ratio in arable crops. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1067498"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"7143","DOI":"10.1002\/ece3.4238","article-title":"Root:shoot ratio in developing seedlings: How seedlings change their allocation in response to seed mass and ambient nutrient supply","volume":"8","author":"Herben","year":"2018","journal-title":"Ecol. Evol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/S0304-4238(00)00152-7","article-title":"Effects of mineral nutrient deficiencies on leaf development, visual symptoms and shoot-root ratio of Spathiphyllum","volume":"86","author":"Yeh","year":"2000","journal-title":"Sci. Horti."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1016\/j.jplph.2007.11.002","article-title":"Boron deficiency decreases growth and photosynthesis, and increases starch and hexoses in leaves of citrus seedlings","volume":"165","author":"Han","year":"2008","journal-title":"J. Plant Physiol."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Meng, X., Chen, W.W., Wang, Y.Y., Huang, Z.R., Ye, X., Chen, L.S., and Yang, L.T. (2021). Effects of phosphorus deficiency on the absorption of mineral nutrients, photosynthetic system performance and antioxidant metabolism in Citrus grandis. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0246944"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1237","DOI":"10.1007\/s00468-012-0699-2","article-title":"Physiological impacts of magnesium-deficiency in citrus seedlings: Photosynthesis, antioxidant system and carbohydrates","volume":"26","author":"Yang","year":"2012","journal-title":"Trees Struct. Funct."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1016\/j.aquatox.2018.03.020","article-title":"Identification and molecular characterization of two Cu\/Zn-SODs and Mn-SOD in the marine ciliate Euplotes crassus: Modulation of enzyme activity and transcripts in response to copper and cadmium","volume":"199","author":"Kim","year":"2018","journal-title":"Aquat. Toxicol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"2165","DOI":"10.1093\/jxb\/erg220","article-title":"Both xanthophyll cycle-dependent thermal dissipation and the antioxidant system are up-regulated in grape (Vitis labrusca L cv Concord) leaves in response to N limitation","volume":"54","author":"Chen","year":"2003","journal-title":"J. Exp. Bot."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"15028002","DOI":"10.4238\/gmr.15028002","article-title":"Expression of an alfalfa (Medicago sativa L.) peroxidase gene in transgenic Arabidopsis thaliana enhances resistance to NaCl and H2O2","volume":"15","author":"Teng","year":"2016","journal-title":"Genet. Mol. Res."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Du, H., Huang, Y., Qu, M., Li, Y., Hu, X., Yang, W., Li, H., He, W., Ding, J., and Liu, C. (2020). A maize ZmAT6 gene confers aluminum tolerance via reactive oxygen species scavenging. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.01016"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1078\/0176-1617-01156","article-title":"Aluminium-induced production of oxygen radicals, lipid peroxidation and DNA damage in seedlings of rice (Oryza sativa)","volume":"161","author":"Meriga","year":"2004","journal-title":"J. Plant Physiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1038\/s43016-022-00596-7","article-title":"Nanobiotechnology-based strategies for enhanced crop stress resilience","volume":"3","author":"Zhao","year":"2022","journal-title":"Nat. Food"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1590\/01047760202026022711","article-title":"Negative effects on photosynthesis and chloroplast pigments exposed to lead and aluminum: A meta-analysis","volume":"26","author":"Wolowski","year":"2020","journal-title":"Cerne"},{"key":"ref_78","unstructured":"Marschner, P. (2012). Function of nutrients: Micronutrients. Marschner\u2019s Mineral Nutrition of Higher Plants, Academic Press. [3rd ed.]."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"3233","DOI":"10.1038\/s41598-022-06867-4","article-title":"Understanding calcium functionality by examining growth characteristics and structural aspects in calcium-deficient grapevine","volume":"12","author":"Duan","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1186\/s11658-023-00479-0","article-title":"Multi-omics strategies uncover the molecular mechanisms of nitrogen, phosphorus and potassium deficiency responses in Brassica napus","volume":"28","author":"Fu","year":"2023","journal-title":"Cell. Mol. Biol. Lett."},{"key":"ref_81","first-page":"113","article-title":"Effects of nitrogen deposition and phosphorus addition on photosynthesis and chlorophyll fluorescence characteristics of Chinese fir","volume":"37","author":"Qian","year":"2018","journal-title":"Ecol. Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1093\/aob\/mci145","article-title":"Effects of aluminum on light energy utilization and photoprotective systems in Citrus leaves","volume":"96","author":"Chen","year":"2005","journal-title":"Ann. Bot."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"107884","DOI":"10.1016\/j.carres.2019.107884","article-title":"Glucose: Sweet or bitter effects in plants-a review on current and future perspective","volume":"87","author":"Siddiqui","year":"2020","journal-title":"Carbohydr. Res."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1111\/jipb.12312","article-title":"Glucose alleviates cadmium toxicity by increasing cadmium fixation in root cell wall and sequestration into vacuole in Arabidopsis","volume":"57","author":"Shi","year":"2015","journal-title":"J. Integr. Plant Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"108271","DOI":"10.1016\/j.carres.2021.108271","article-title":"Glucose modulates copper induced changes in photosynthesis, ion uptake, antioxidants and proline in Cucumis sativus plants","volume":"501","author":"Yusuf","year":"2021","journal-title":"Carbohydr. Res."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1215","DOI":"10.1093\/treephys\/tpz042","article-title":"Copper accumulation, subcellular partitioning and physiological and molecular responses in relation to different copper tolerance in apple rootstocks","volume":"39","author":"Wan","year":"2019","journal-title":"Tree Physiol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1023\/A:1026012116709","article-title":"New fluorescence parameters for monitoring photosynthesis in plants","volume":"78","author":"Force","year":"2003","journal-title":"Photosynth. Res."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1863","DOI":"10.1093\/treephys\/28.12.1863","article-title":"Aluminum-induced effects on photosystem II photochemistry in Citrus leaves assessed by the chlorophyll a fluorescence transient","volume":"28","author":"Jiang","year":"2008","journal-title":"Tree Physiol."},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Hao, J., Han, H., Liu, Y., Li, J., Yang, J., Ren, B., and Bai, L. (2003). Phosphorus addition alleviates the inhibition of nitrogen deposition on photosynthesis of Potentilla tanacetifolia. Front. Environ. Sci., 11.","DOI":"10.3389\/fenvs.2023.1099203"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1007\/s11120-017-0467-7","article-title":"Chlorophyll fluorescence as a tool for nutrient status identification in rapeseed plants","volume":"136","author":"Kalaji","year":"2018","journal-title":"Photosynth. Res."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/0076-6879(87)48036-1","article-title":"Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes","volume":"148","author":"Lichtenthaler","year":"1987","journal-title":"Method. Enzymol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1097\/00010694-196310000-00002","article-title":"Effect of initial pH, phosphate, and silicate on the determination of aluminum with aluminon","volume":"96","author":"Hsu","year":"1963","journal-title":"Soil Sci."},{"key":"ref_93","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_94","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1104\/pp.59.2.309","article-title":"Superoxide dismutases I. Occurrence in higher plants","volume":"59","author":"Giannopolitis","year":"1977","journal-title":"Plant Physiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1104\/pp.60.3.379","article-title":"Enzymic assay of 10\u22127 to 10\u221214 moles of sucrose in plant tissues","volume":"60","author":"Jones","year":"1977","journal-title":"Plant Physiol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1042\/bj0570508","article-title":"The estimation of carbohydrates in plant extracts by anthrone","volume":"57","author":"Yemm","year":"1954","journal-title":"Biochem. J."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"1564","DOI":"10.1093\/treephys\/tpx067","article-title":"Magnesium-deficiency-induced alterations of gas exchange, major metabolites and key enzymes differ among roots, and lower and upper leaves of Citrus sinensis seedlings","volume":"37","author":"Li","year":"2017","journal-title":"Tree Physiol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","article-title":"A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding","volume":"72","author":"Bradford","year":"1976","journal-title":"Anal. Biochem."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/15\/2370\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T06:46:30Z","timestamp":1785739590000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/15\/2370"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,31]]},"references-count":98,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["plants15152370"],"URL":"https:\/\/doi.org\/10.3390\/plants15152370","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,31]]}}}