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

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Paper 3photosynthesis of Lobelia leaves at all organicenrichments whereas TN, chlorophyll andphotosynthesis of Littorella leaves are notsignificantly reduced by any organic enrichmentand only TP is reduced at 0.8 and 1.6 % organicmatter.TP declined more significantly than TNin the leaves by organic sediment enrichmentand TP in Lobelia in several instances droppedbelow the minimum average threshold ofsubmerged macrophytes to sustain growth (0.08% P; Colman et al. 1987, Demars and Edwards2007) both in laboratory and field experiments.3-While PO 4 remained below analyticaldetection (0.2 µM) in the pore-water oforganically enriched sediments due to strongadsorption, NH +4 reached high concentrationsand can be supplied to the plants by passivediffusion (Marchner 1986). P-supply to roots viaVA-fungi is also believed to be impeded bysediment anoxia (Wigand et al. 1998) andinfection of new roots is grossly reduced inorganically enriched anoxic sediments (Møller,Kjøller & Sand-Jensen, pers. comm.).Our results, therefore, imply that low gaspermeability of leaf surfaces is essential for theexcellent ability of Lobelia to thrive onextremely low-organic, nutrient-poor mineralsediments by ensuring an intimate O 2 and CO 2exchange between plant and sediment andpermitting the same leaves to function in air andunder water. In contrast, even small enrichmentof sediments with labile organic matter led toprogressive problems for Lobelia’s performanceand survival by inducing sediment and plantanoxia, impeding P uptake and, finally, stoppingvirtually all photosynthesis and growth.Littorella by possessing gas permeable leafsurfaces is less likely to experience anoxia andmandatory nutrient supply, photosynthesis andgrowth and can much better cope and evenprofit from sediment enrichment. The maindisadvantage for Littorella is that it must investin new leaves upon air exposure or submergenceputting an extra burden on the species undervery nutrient-poor conditions but permitting it toadapt and grow faster under richer conditions.Thus, a distinct anatomical trait, i.e., leaf gaspermeability appears to have heavyconsequences for the fundamental and realizedniche of the two species and, therefore, for theirdistribution and survival in an increasinglyeutrophied environment.AcknowledgementsWe thank Tim D. Colmer for commenting on themanuscript and Birgit Kjøller and Annette Adamssonfor technical assistance. This project was funded bya grand from the Willum Kann RasmussenFoundation to Center of Lake Restoration (CLEAR).ReferencesArmstrong W, Cousins D, Armstrong J, Turner DW,Beckett PM 2000. Oxygen distribution in wetland plantroots and permeability barriers to gas-exchange with therhizonsphere: a microelectrode and modeling study withPhragmites australis. Annals of Botany 54: 177-198.Arts GHP 2002. Deterioration of Atlantic softwatermacrophyte communities by acidification, eutrophicationand alkalinisation. Aquatic Botany 73: 273-393.Christensen PB, Revsbech NP, Sand-Jensen K 1994.Microsensor analysis of oxygen in the rhizosphere of theaquatic macrophyte Littorella uniflora (L.) Aschers. PlantPhysiology 105: 1174-1179.Colman JA, Sorsa K, Hofmann JP, Smith CS,Andrews JH 1987. Yield-derived and photosynthesisderivedcritical concentrations of tissue phosphorus andtheir significance for growth of Eurasian Water Milfoil,Myriophyllum spicatum L. Aquatic Botany 29: 111-122.66
Paper 3Colmer TD 2003. Long-distance transport of gases inplants: a perspective on internal aeration and radialoxygen loss from roots. Plant, Cell and Environment 26:17-36.Colmer TD, Flowers TJ 2008. Flooding tolerance inhalophytes. New Phytologist 179: 964-974.Demars BOL, Edwards AC 2007. Tissue nutrientconcentrations in freshwater aquatic macrophytes: highinter-taxon differences and low phenotypic response tonutrient supply. Freshwater Biology 52: 2073-2086.Gerloff GC, Krombholz PH 1966. Tissue analysis as ameasure of nutrient availability for the growth ofangiosperm aquatic plants. Limnology and Oceanography11: 529-537.Gibbs J, Greenway H 2003. Mechanisms of anoxiatolerance in plants. I. Growth, survival and anaerobiccatabolism. Functional Plant Biology 30: 1-47.Greenway H, Gibbs J 2003. Mechanisms of anoxiatolerance in plants. II- Energy requirements formaintenance and energy distribution to essentialprocesses. Functional Plant Biology 30: 999-1036.Madsen TV 1985. A community of submerged aquaticCAM plants in Lake Kalgaard, Denmark. Aquatic Botany23: 97-108.Marschner H. 1986. Mineral nutrition of higher plants.Academic Press, London.Nielsen SL, Gacia E, Sand-Jensen K 1991. Land plantsof amphibious Littorella uniflora (L.) Aschers. maintainutilization of CO 2 from sediments. Oecologia 88: 258-263.Møller CL, Sand-Jensen K 2008. Iron plaques improvethe oxygen supply to root meristems of the freshwaterplant, Lobelia dortmanna. New Phytologist 179: 848-856.Møller CL, Sand-Jensen K 2011. High sensitivity ofLobelia dortmanna to sediment oxygen depletionfollowing organic enrichment. New Phytologist 190: 320-331.Pedersen O, Andersen T, Ekejima K, Hossain MZ,Andersen FØ 2006. A multidisciplinary approach tounderstanding the recent and historical occurrence of thefreshwater plant Littorella uniflora. Freshwater Biology51: 865-873.Pedersen O, Sand-Jensen K, Revsbech NP 1995. Dielpulses of O 2 and CO 2 in sandy sediments inhabited byLobelia dortmanna. Ecology 76: 1536-1545.Pedersen O, Sand-Jensen K 1992. Adaptations ofsubmerged Lobelia dortmanna to aerial life form:morphology, carbon sources and oxygen dynamics. Oikos65: 89-96.Raun AL, Borum J, Sand-Jensen K 2010. Influence ofsediment organic enrichment and water alkalinity ongrowth of aquatic isoetid and elodeid plants. FreshwaterBiology 55: 1891-1904.Sand-Jensen K 1987. Environmental control ofbicarbonate use among freshwater and marinemacrophytes. In RM Crawford (ed), Ecology andPhysiology of Intertidal plants, pp. 99-112. Blackwell,Oxford.Sand-Jensen K, Borum J 1991. Interactions amongphytoplankton, periphyton and macrophytes in temperatefreshwaters. Aquatic Botany 41: 137-175.Sand-Jensen K, Borum J, Binzer T 2005. Oxygen stressand reduced growth of Lobelia dortmanna in sandy lakesediments subject to organic enrichment. FreshwaterBiology 50: 1034-1048.Sand-Jensen K, Riis T, Vestergaard O, Larsen SE2000. Macrophyte decline in Danish lakes and streamsover the past 100 years. Journal of Ecology 88: 1030-1040.Sand-Jensen K, Prahl C 1982. Oxygen exchange withthe lacunae and across leaves and roots of the submergedvascular macrophyte, Lobelia dortmanna L. NewPhytologist 91: 103-120.Sand-Jensen K, Prahl C, Stokholm H 1982. Oxygenrelease from roots of submerged aquatic macrophytes.Oikos 38: 349-354.Sand-Jensen K, Søndergaard M 1978. Growth andproduction of isoetids in oligotrophic Lake Kalgaard,Denmark. Internationale Vereiningung füt theoretischeund angewandte Limnologie, Verhandlungen 9: 659-666.Sand-Jensen K, Søndergaard M 1979. Distribution andquantitative development of aquatic macrophytes inrelation to sediment characteristics in oligotrophic LakeKalgaard, Denmark. Freshwater Biology 9: 1-11.Smolders AJP, Lucassen ECHE, Roelofs JGM 2002.The isoetid environment: biogeochemistry and threats.Aquatic Botany 73: 325-350.Stelzer D, Schneider S, Melzer A 2005. Macrophytebasedassessment of lakes – a contribution for theimplementation of the European Water Framework67
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Oxygen dynamics and plant-sedimenti
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PrefaceThe content of this thesis i
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AbstractAbstract● Isoetids occupy
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IntroductionIntroductionOligotrophi
Page 15 and 16: Introductionwater (Wigand et al. 19
Page 17 and 18: IntroductionSand-Jensen K, Prahl C,
Page 19 and 20: Thesis summaryThesis summaryAdditio
Page 21 and 22: Thesis summaryFig. 4. Chlorophyll c
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Page 27: Paper 1Oxygen, carbon and iron dyna
Page 30 and 31: Paper 1extensive sediment volume by
Page 32 and 33: Paper 1method of Andersen (1976) an
Page 35 and 36: Paper 1Table 3. Retention of added
Page 37 and 38: Paper 1recalcitrant organic pools a
Page 39: Paper 2High sensitivity of Lobelia
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Page 52 and 53: NewPaper 2Phytologist Research 331c
Page 55 and 56: Paper 3Higher gas permeability of l
Page 57 and 58: Paper 3Here, we perform a parallel
Page 59 and 60: Paper 3Fig. 1. O 2 penetration dept
Page 61 and 62: Paper 3Table 2. O 2 flux across lea
Page 63 and 64: Paper 3Fig. 4. Leaf chlorophyll of
Page 65: Paper 3high internal concentrations
Page 69: Paper 4Organic enrichment of sedime
Page 72 and 73: Paper 4Surveys on aquatic plants re
Page 74 and 75: Paper 4ensure mixing and air satura
Page 76 and 77: Paper 4ResultsFig 1. O 2 -penetrati
Page 78 and 79: Paper 4Table 2. Individual leaf bio
Page 80 and 81: Paper 4Fig. 4. Root density (○) a
Page 82 and 83: Paper 4hours every night although l
Page 84 and 85: Paper 4Møller CL, Sand-Jensen K. 2
Page 87 and 88: [Plant Signaling & Behavior 3:10, 8
Page 89: Outstanding Lobelia dortmanna in ir
Page 93 and 94: Planterødders overlevelse ivanddæ
Page 95 and 96: Paper 6Figur 3. Aflejringer af lign
Page 97: Paper 6Vi har sat planterødders ve
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