620bbb19d7b3129863880a8e937692ce plants-15-01877.pdf 1da958c9872c6358fc6d7a02a43893d2c7804ab5 plants-15-01877.pdf bc308e6a61ee29cca3267858a763728cc8ed920d50c6994d1ac7cb524e85cc05 plants-15-01877.pdf Title: Bryophytes as Strong Aluminum Accumulators in Acidic Soils: Cell-Wall Binding and Physiological Tolerance Mechanisms Subject: Bryophytes are key components of acid–soil 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 g g-1 DW), with some reaching values exceeding 28,000 g g-1 DW, confirming their strong accumulation capacity. After Al, iron was the most abundantly accumulated metal (1430–22,800 g g-1 DW), followed by manganese (100–3100 g g-1 DW). The sampled lichen species accumulated Al at concentrations between 1063 and 9154 g g-1 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—Barbula unguiculata, Palamocladium euchloron, and Hypnum cupressiforme—were acclimated to laboratory conditions and treated with two Al levels (without or with 150 M 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—phenolics, flavonoids, and carboxylic acids (tartaric, oxalic, malic, and citric acids)—were 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. Keywords: aluminum accumulation; aluminum sequestration; callose; carboxylic acids; cell-wall binding; liverworts; mosses; pectin Author: Roghieh Hajiboland, Aiuob Moradi, Hedieh Majmoueh-Koub, Roser Tolrà, Ana Paravinja, Milos Stanojevic, Miroslav Nikolic and Charlotte Poschenrieder Creator: LaTeX with hyperref Producer: pdfTeX-1.40.25; modified using OpenPDF 1.4.2 CreationDate: Wed Jun 17 08:33:27 2026 CEST ModDate: Wed Jun 17 08:44:37 2026 CEST Custom Metadata: yes Metadata Stream: no Tagged: no UserProperties: no Suspects: no Form: none JavaScript: no Pages: 28 Encrypted: no Page size: 595.276 x 841.89 pts (A4) Page rot: 0 File size: 8069610 bytes Optimized: no PDF version: 1.7 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- XPULKE+URWPalladioL-Ital Type 1 Custom yes yes yes 206 0 WNWTFR+VnURWPalladioL-Bold Type 1 Custom yes yes yes 207 0 CAMXTK+EURM10 Type 1 Builtin yes yes yes 208 0 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LOJIKP+PalatinoLinotype TrueType WinAnsi yes yes no 1020 0 LOJIKP+PalatinoLinotype TrueType WinAnsi yes yes no 1207 0 HQBQWV+VnURWPalladioL Type 1 Custom yes yes yes 1585 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-09-06 03:56:19 CEST RepresentationInformation: plants-15-01877.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-09-05 08:08:30 CEST Size: 8069610 Format: PDF Version: 1.7 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf PDFMetadata: Objects: 1727 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: true CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseNone Outlines: Item: Title: Introduction Item: Title: Results Children: Item: Title: Accumulation of Al in Collected Bryophyte Specimens Item: Title: Accumulation of Al in Other Non-Seed Plants Collected Item: Title: Accumulation of Other Elements in Collected Bryophyte Specimens and Its Correlation with Al Item: Title: Aluminum Concentration, Photosynthetic Pigments, Osmolytes, and Callose in the Leafy Shoots of Field-Collected, Laboratory-Acclimated Bryophyte Species Item: Title: Accumulation and Localization of Al in Field-Collected, Laboratory-Acclimated Bryophyte Species Item: Title: Antioxidant Defense Responses of Al-Treated Mosses Item: Title: Activity of Phenolic-Metabolizing Enzymes and Concentrations of Al-Chelating Compounds in Al-Treated Mosses Item: Title: C2 Metabolism Intermediates and Formic Acid in Al-Treated Mosses Item: Title: FTIR Analysis and Cell-Wall Characteristics in Al-Treated Mosses Item: Title: Discussion Children: Item: Title: Widespread and Substantial Al Accumulation in Bryophytes Item: Title: Acidic Soil Chemistry Drives Coordinated Metal Accumulation in Bryophytes Item: Title: Oxidative Stress and Osmotic Adjustment Responses to Al Exposure in Bryophytes Item: Title: Phenolic Compounds Indicate Metabolic Adjustments and Detoxification Responses Item: Title: Carboxylates in Species-Specific Metabolic Adjustment and Al Detoxification Item: Title: Callose Deposition and Its Limited Role in Al Tolerance Item: Title: Evidence for Al Immobilization and Cell-Wall-Based Al Tolerance Item: Title: Interspecific Differences in Al Accumulation and the Expression of Al Detoxification Mechanisms Item: Title: Ecological Implications of Al Accumulation Item: Title: Materials and Methods Children: Item: Title: Study Area, Collection of Specimens, and Identification Item: Title: Elemental Analyses of Collected Specimens Item: Title: Culture of Bryophytes Under Growth Chamber Conditions Item: Title: Quantification of Al, Pigments, Carbohydrates, and Proline in Laboratory-Cultivated Mosses Item: Title: Histochemical Analyses of Laboratory-Cultivated Mosses Item: Title: Activity of Antioxidant Enzymes and Concentration of Oxidant Metabolites Item: Title: Activity of Phenolic-Metabolizing Enzymes and Concentration of Phenolic Compounds Item: Title: Analysis of Carboxylic Acids Item: Title: FTIR Analysis Item: Title: Statistical Analyses Item: Title: Conclusions Item: Title: References Info: Title: Bryophytes as Strong Aluminum Accumulators in Acidic Soils: Cell-Wall Binding and Physiological Tolerance Mechanisms Author: Roghieh Hajiboland, Aiuob Moradi, Hedieh Majmoueh-Koub, Roser Tolrà, Ana Paravinja, Milos Stanojevic, Miroslav Nikolic and Charlotte Poschenrieder Subject: Bryophytes are key components of acid–soil 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 g g-1 DW), with some reaching values exceeding 28,000 g g-1 DW, confirming their strong accumulation capacity. After Al, iron was the most abundantly accumulated metal (1430–22,800 g g-1 DW), followed by manganese (100–3100 g g-1 DW). The sampled lichen species accumulated Al at concentrations between 1063 and 9154 g g-1 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—Barbula unguiculata, Palamocladium euchloron, and Hypnum cupressiforme—were acclimated to laboratory conditions and treated with two Al levels (without or with 150 M 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—phenolics, flavonoids, and carboxylic acids (tartaric, oxalic, malic, and citric acids)—were 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. 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