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

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Paper 3Higher gas permeability of leaves provides greater tolerance of Littorella uniflorathan Lobelia dortmanna to <strong>sediment</strong> organic enrichmentClaus L<strong>in</strong>dskov Møller <strong>and</strong> Kaj S<strong>and</strong>-JensenFreshwater Biological Laboratory, Biological Institute, University of Copenhagen, Hels<strong>in</strong>gørsgade 51, DK-3400 Hillerød,DenmarkKey words: Permeability,<strong>isoetid</strong>s, eutrophication,biogeochemistry, anoxia,oxygen, stress.Summary● Lobelia dortmanna <strong>and</strong> Littorella uniflora are prom<strong>in</strong>ent <strong>in</strong> nutrientpoor,softwater lakes because of efficient root exchange of CO 2 <strong>and</strong> O 2 .We hypothesize that higher gas permeability of Littorella than Lobelialeaves ensures O 2 uptake from water <strong>and</strong> greater tolerance to organicenrichment <strong>and</strong> anoxia of <strong>sediment</strong>s.● We studied O 2 <strong>dynamics</strong> <strong>in</strong> <strong>plant</strong>s <strong>and</strong> <strong>sediment</strong>s <strong>and</strong> leaf photosynthesisupon organic <strong>sediment</strong> enrichment <strong>in</strong> long-term laboratory <strong>and</strong> fieldstudies.● Addition of organic matter triggered O 2 depletion <strong>and</strong> accumulation ofNH 4 + , Fe 2+ <strong>and</strong> CO 2 <strong>in</strong> <strong>sediment</strong>s. O 2 <strong>in</strong> Lobelia’s leaf lacunae fluctuatedfrom air saturation <strong>in</strong> the light to anoxia late at night on natural <strong>sediment</strong>s,but organic enrichment prolonged night anoxia. Littorella’s leaf surfaceswere 13-16 times more permeable <strong>and</strong> O 2 <strong>in</strong> the lacunae rema<strong>in</strong>ed above10 kPa on anoxic <strong>sediment</strong>s due to uptake from the water. Leaf N, P <strong>and</strong>photosynthesis dropped to critically low levels under prolonged <strong>sediment</strong>anoxia for Lobelia but not for Littorella.● High gas permeability of Littorella leaves improves performance <strong>and</strong>survival on organically enriched, anoxic <strong>sediment</strong>s compared with Lobelia.Littorella changes between gas impermeable aerial leaves <strong>and</strong> permeableaquatic leaves whereas Lobelia uses the same impermeable leaves <strong>in</strong> air<strong>and</strong> water which is cost-effective <strong>in</strong> ultra-oligotrophic environments.IntroductionOligotrophic, softwater lakes <strong>in</strong> Europe <strong>and</strong>North America are <strong>in</strong>habited by a taxonomicallydiverse group of small <strong>isoetid</strong> species all hav<strong>in</strong>gleaf rosettes, large root systems <strong>and</strong> extensiveaerenchyma as adaptations to acquire CO 2 <strong>and</strong>nutrients from <strong>sediment</strong>s (Smolders et al. 2002).The <strong>isoetid</strong>s are key species <strong>in</strong> <strong>in</strong>ternationalevaluations of prist<strong>in</strong>e nature quality such asEU’s Habitat Directive <strong>and</strong> Water FrameworkDirective (Stelzer et al. 2005) <strong>and</strong> oligotrophic,softwater lakes are named Lobelia-lakes afterthe conspicuous flower<strong>in</strong>g species, Lobeliadortmanna. Despite the same growth form <strong>and</strong>requirement of relatively unpolluted habitats ofall <strong>isoetid</strong>s, the species vary substantially <strong>in</strong>anatomy <strong>and</strong> physiology which may account fordifferences <strong>in</strong> their distributions (S<strong>and</strong>-Jensen &Søndergaard 1979). For example, Littorellauniflora has a wider ecological range <strong>and</strong>tolerates greater eutrophication than Lobeliadortmanna which has gone ext<strong>in</strong>ct <strong>in</strong> many NWEuropean lakes <strong>and</strong> it is now conf<strong>in</strong>ed to a fewprist<strong>in</strong>e oligotrophic lakes (Arts et al. 1989;S<strong>and</strong>-Jensen et al. 2000; Pedersen et al. 2006).We here test if Littorella is more tolerant thanLobelia to organic enrichment <strong>and</strong> anoxia of55

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