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

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Paper 1Table 3. Retention of added C, N <strong>and</strong> P to <strong>sediment</strong>s at term<strong>in</strong>ation of the c. 200-days longexperiments with Lobelia (Lob) <strong>and</strong> Littorella (Lit).C (mol m -2 ) TN (mol m -2 ) TP (mol m -2 )Treatmen Added LeftLobLeftLitAdded LeftLobLeftLitAdded LeftLobLeftLit0 % 0 0 0 0 0 0 0 0 00.1 % 5.8 4.6 -2.9 0.24 0.16 0.15 0.013 0.017 0.0080.2 % 11.6 3.5 5.8 0.49 0.24 0.60 0.026 0.024 0.0480.4 % 23.1 13.3 6.9 0.97 1.00 -0.10 0.051 0.047 0.0320.8 % 46.2 26.6 13.3 1.94 1.31 1.39 0.103 0.095 0.0631.6 % 92.4 58.9 39.3 3.89 4.02 1.98 0.206 0.121 0.149the two species, while retention wassignificantly smaller for C (44±35%), <strong>and</strong> alsotended to be lower for N (77±26%) than P (ttest;Table 3). Retentions were not significantlydifferent between Lobelia <strong>and</strong> Littorella.DiscussionIn natural low-organic <strong>sediment</strong>s of low O 2consumption rates, high O 2 release frompermeable roots of <strong>isoetid</strong>s builds-up O 2concentrations above air saturation <strong>in</strong> the porewater<strong>in</strong> the light (Møller <strong>and</strong> S<strong>and</strong>-Jensen2011a,b) <strong>and</strong> O 2 penetrates to more than 40 mmdepth <strong>in</strong> the <strong>sediment</strong>s. Lobelia has leaf surfacesof low gas permeability <strong>and</strong> releases mostphotosynthetic O 2 <strong>in</strong> the light from the roots tothe <strong>sediment</strong> <strong>and</strong> takes up O 2 by the roots <strong>in</strong> thedark until the pore-water <strong>and</strong> the <strong>plant</strong> tissueturn anoxic (Møller <strong>and</strong> S<strong>and</strong>-Jensen 2011a).Littorella’s leaf surfaces have 12-14 timeshigher permeability than Lobelia’s (Møller <strong>and</strong>S<strong>and</strong>-Jensen 2011b) <strong>and</strong> releases about 30% ofphotosynthetic O 2 from the roots <strong>in</strong> the light(S<strong>and</strong>-Jensen et al. 1982), while O 2 uptake fromlake water <strong>in</strong> the dark ensures O 2 concentrations<strong>in</strong> the leaf lacunae above 5-10 Pa (Møller <strong>and</strong>S<strong>and</strong>-Jensen 2011b), downstream transport toroots <strong>and</strong> release to <strong>sediment</strong>s at about 20-40%of the rate <strong>in</strong> the light (Christensen et al. 1994).The more rapid recovery of O 2 penetrationdepths <strong>and</strong> disappearance of Fe 2+ after 0.1-0.4%organic enrichment of Littorella but not Lobelia<strong>sediment</strong>s <strong>and</strong> the small efflux <strong>and</strong>accumulation of Fe 2+ relative to DIC ofLittorella <strong>sediment</strong>s suggest that O 2 supply ishigher to Littorella <strong>sediment</strong>s than to Lobelia<strong>sediment</strong>s. Three times higher leaf biomass, 1.3-1.5 times higher mass specific photosynthesis(Møller <strong>and</strong> S<strong>and</strong>-Jensen 2011b) <strong>and</strong>appreciable dark release of Littorella(Christensen et al. 1994) can account for anestimated higher O 2 release to <strong>sediment</strong>s withLittorella (20-40 mmol m -2 d -1 ) than Lobelia(10-20 mmol m -2 d -1 ).Well oxygenated conditions of<strong>sediment</strong>s with no or low organic enrichment(0.1%) ensure that aerobic respiration is theprom<strong>in</strong>ent respiration mode. O 2 depth profiles <strong>in</strong>the light show one peak a few mm below the<strong>sediment</strong> surface due to photosynthesis ofbenthic microalgae <strong>and</strong> a second peak at 20 mmdue to O 2 release from <strong>isoetid</strong> roots (Pedersen etal. 1995, S<strong>and</strong>-Jensen et al. 2005). High O 2concentrations permit conversion of NH +4 toNO - 3 that can drive denitrification dur<strong>in</strong>g anoxia<strong>in</strong> two zones below a surface <strong>and</strong> a deeper zoneof maximum O 2 concentrations at depth<strong>in</strong>tegratedrates of 500-700 µmol m -2 d -1 <strong>in</strong> fieldmeasurements on Littorella <strong>and</strong> laboratoryexperiments with Lobelia (Christensen <strong>and</strong>Sørensen 1986, Risgaard-Petersen <strong>and</strong> Jensen1997). The experiments with Lobelia showedthat denitrification was stimulated by a factorgreater than six when compared to bare<strong>sediment</strong>s <strong>and</strong> this enhanced activity was due to+root mediated oxic stimulation of both NH 4-release <strong>and</strong> NO 3 formation driv<strong>in</strong>g35

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