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Oxygen dynamics and plant-sediment interactions in isoetid ...

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NewPaper 2Phytologist Research 323Värsjö as used for collection of <strong>sediment</strong> turf for laboratoryexperiments to make sure that patterns observed <strong>in</strong> laboratoryexperiments resembled natural <strong>sediment</strong> <strong>and</strong> <strong>plant</strong>responses <strong>in</strong> the field.A long-term organic enrichment experiment was madeover 80 d <strong>in</strong> the field (10 July–1 October) <strong>in</strong> which <strong>sediment</strong>chemistry <strong>and</strong> <strong>plant</strong> performance were comparedbetween triplicate controls <strong>and</strong> treatments (20 · 20 cm perplot) enriched with organic matter to 0.8% DW asdescribed for the laboratory experiment. At the end of theexperiment, pore-water concentrations <strong>and</strong> <strong>sediment</strong> compositionwere measured on <strong>sediment</strong> cores (5 cm <strong>in</strong>diameter, 10 cm <strong>in</strong> depth) taken from each plot <strong>and</strong>brought back to the laboratory <strong>in</strong> Perspex tubes with rubberstoppers prevent<strong>in</strong>g disturbance of <strong>sediment</strong> <strong>and</strong> porewater. Cores were stored <strong>in</strong> the laboratory at 15°C <strong>in</strong>a12 : 12 h light : dark cycle for 1–4 d before pore water wasextracted <strong>and</strong> analysed as already described. Likewise, netphotosynthesis was measured <strong>in</strong> the laboratory <strong>in</strong>cubator onleaf samples from each plot. Leaf samples derived fromharvested field <strong>plant</strong>s were transported to the laboratory <strong>in</strong>sealed moist plastic bags <strong>and</strong> stored at 8°C <strong>and</strong> used forexperiments with<strong>in</strong> 2 d.O 2 <strong>dynamics</strong> was measured cont<strong>in</strong>uously over a full 24 hlight–dark cycle <strong>in</strong> the field <strong>in</strong> late August <strong>in</strong> duplicated experimentsby <strong>in</strong>sert<strong>in</strong>g microelectrodes <strong>in</strong>to leaf lacunae of two<strong>plant</strong>s <strong>and</strong> <strong>in</strong>sert<strong>in</strong>g m<strong>in</strong>i-electrodes at 1 cm depth <strong>in</strong> the<strong>sediment</strong> next to these two <strong>plant</strong>s us<strong>in</strong>g the same type of equipmentas for laboratory experiments. HOBO data loggers wereused to measure water temperature <strong>and</strong> irradiance (OnsetComputer Corporation, Bourne, Massachusetts, USA).Data treatment <strong>and</strong> statistical analysisData were processed <strong>in</strong> Excel 2007 <strong>and</strong> statistical analyses<strong>and</strong> graphs were made <strong>in</strong> Graph Pad Prism 5. Data arepresented as means ± SD. P < 0.05 was considered significant.The <strong>in</strong> situ experiment was a block design <strong>in</strong> triplicatesuitable for t-test analysis, whereas the laboratory experimentwas a gradient study with six amounts of added organicmatter to s<strong>in</strong>gle <strong>sediment</strong> turfs <strong>in</strong>habited by 20 <strong>plant</strong>s(pseudo-replicates) suited for regression ⁄ correlation analysis.Values from the laboratory experiment are presented asmeans ± SD of usually three replicates from each <strong>sediment</strong> turf.ResultsSediment processes <strong>and</strong> chemistryOrganic enrichment of <strong>sediment</strong>s <strong>in</strong> long-term laboratoryexperiments stimulated O 2 use for degradation of organicmatter <strong>and</strong> reduced O 2 depth penetration (Fig. 1). O 2 penetrationdepth exceeded 40 mm <strong>in</strong> control <strong>sediment</strong>s butdecl<strong>in</strong>ed to < 3 mm after 25 d <strong>in</strong> <strong>sediment</strong>s fertilized withFig. 1 Depth of O 2 penetration <strong>in</strong> Lobelia <strong>sediment</strong>s as a functionof time after addition of different amounts of labile organic matter(per <strong>sediment</strong> DW) (open circles, control; closed circles, 0.1%; opensquares, 0.2%; closed squares, 0.4%; open diamonds, 0.8%; closeddiamonds, 1.6%). Measurements were made 10–12 h <strong>in</strong>to the 12 hlight period. Measurements could not extend deeper than 40 mm<strong>in</strong>to the <strong>sediment</strong>, when O 2 penetration <strong>in</strong> control <strong>sediment</strong>s wasdescribed as > 40 mm. Values are mean of two (0–170 d) <strong>and</strong> three(194 d) measurements ± SD.‡ 0.4% organic matter. In <strong>sediment</strong>s receiv<strong>in</strong>g only 0.1 <strong>and</strong>0.2% organic matter, O 2 penetration depth had onlydecl<strong>in</strong>ed to 21 <strong>and</strong> 10 mm, respectively, 25 d after fertilization.As degradation of organic matter progressed overtime, O 2 gradually penetrated deeper <strong>in</strong>to the <strong>sediment</strong>s.However, only at the lowest organic dose did O 2 penetrationrecover to the same value (> 40 mm) as <strong>in</strong> the control<strong>sediment</strong> with<strong>in</strong> the 195 d of experiments. At the end ofthe experiment, O 2 penetration depth was significantly<strong>and</strong> negatively correlated to the magnitude of addition(Spearman’s r, P < 0.001). Thus, organic enrichment hadprofound <strong>and</strong> long-last<strong>in</strong>g effects on O 2 availability <strong>and</strong>decomposition processes.This persistent enrichment effect was evident <strong>in</strong> <strong>sediment</strong>chemistry even at the lowest enrichment after 194 d <strong>in</strong>the laboratory experiment (Table 1). Across the gradient,organic content, TN <strong>and</strong> TP <strong>in</strong>creased two- to threefold<strong>and</strong> they were significantly correlated to the amount oforganic matter added (l<strong>in</strong>ear regression, P < 0.0001), whilewater content <strong>and</strong> TFe did not change significantly.Approximately 14.5 mg organic matter, 368 lg TN <strong>and</strong>43 lg TP (all g –1 dry <strong>sediment</strong>) were added with the highest1.6% organic enrichment <strong>and</strong> the treated <strong>sediment</strong> stillconta<strong>in</strong>ed an extra 10.2 mg organic matter, 382 lg TN<strong>and</strong> 25.5 lg TP relative to the control <strong>sediment</strong> after194 d. Likewise, the 0.4% organic treatment received anadditional 3.6 mg organic matter, 92 lg TN <strong>and</strong> 9.8 lgTP per g dry <strong>sediment</strong> <strong>and</strong> still conta<strong>in</strong>ed extra 2.3 mgorganic matter, 96 lg TN <strong>and</strong> 9.8 lg TP after 194 d,stress<strong>in</strong>g that organic matter was lost by degradation butmost N <strong>and</strong> P rema<strong>in</strong>ed <strong>in</strong> the <strong>sediment</strong>.Ó 2010 The AuthorsNew Phytologist Ó 2010 New Phytologist Trust44New Phytologist (2011) 190: 320–331www.newphytologist.com

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