Paper 4hours every night although low O 2 tension hasbeen shown to prevent AMF development (LeTacon et al. 1983).Plant growth, stress <strong>and</strong> nutritionPlants were clearly stressed by high organicaddition <strong>and</strong> this was always coupled with lowerAMF colonization. In experiments with <strong>in</strong>tact<strong>sediment</strong> turfs, P levels <strong>in</strong> leaves tended todecrease at the same organic addition (0.4%)caus<strong>in</strong>g AMF colonization to drop (Fig. 2; Table3). In contrast, Littorella was able to grow <strong>and</strong>ma<strong>in</strong>ta<strong>in</strong> unaltered nutrient <strong>and</strong> chlorophylllevels <strong>and</strong> biomasses with very low AMFcolonization at low organic enrichments (0.2%<strong>in</strong> the colonization experiment) as <strong>plant</strong>s withhigh AMF colonization <strong>in</strong> control <strong>sediment</strong>s(Table 1). There is, therefore, no clear evidenceof AMF be<strong>in</strong>g responsible for the observedstress. It has been shown for the <strong>isoetid</strong> Isoetesalp<strong>in</strong>us that root anoxia stops translocation ofphotosynthates from leaves to roots (Sorrell2004) which can lead to root malfunction <strong>and</strong>decreased <strong>plant</strong> nutrition (Møller & S<strong>and</strong>-Jensen2011) <strong>and</strong> this is the most plausible reason forthe observed stress. Furthermore, under these<strong>sediment</strong> conditions shad<strong>in</strong>g of the <strong>isoetid</strong>s byfaster grow<strong>in</strong>g <strong>plant</strong>s or filamentous algae islikely to occur (S<strong>and</strong>-Jensen 2000; Arts 2002).It is, therefore, more reasonable to regard AMFas an advantage under very nutrient-poorconditions whereas transport of photosynthatesto AMF under more nutrient-rich <strong>sediment</strong>conditions can be spared because <strong>plant</strong>s cansusta<strong>in</strong> sufficient nutrient uptake without thepresence of AMF (Smith & Read 2008).This <strong>in</strong>vestigation showed that <strong>isoetid</strong>swith high AMF colonization have extensivehyphal networks <strong>in</strong>creas<strong>in</strong>g <strong>sediment</strong> contact<strong>and</strong> surface area for nutrient uptake <strong>in</strong> nutrientpoor<strong>sediment</strong>s. AMF colonization <strong>in</strong> <strong>plant</strong>sfrom mixed populations <strong>in</strong> Värsjö was similar tothat <strong>in</strong> long-term laboratory experiments withmono-specific populations <strong>in</strong> Littorella <strong>and</strong>Lobelia turfs collected <strong>in</strong> the field. Littorellahad higher root colonization <strong>and</strong> a higher AMFcolonization per leaf weight than Lobelia <strong>and</strong> isknown to have two times higher growth rateunder field conditions (S<strong>and</strong>-Jensen &Søndergaard 1978; Bosten & Adams 1989).Thus, Littorella needs a higher nutrient uptakeper biomass than Lobelia. Higher colonizationof Littorella roots be<strong>in</strong>g replaced more rapidlythan Lobelia roots requires higher colonizationrates. This higher colonization rate of AMFcould be supported by higher downward O 2supply to Littorella roots <strong>and</strong> <strong>sediment</strong>s dur<strong>in</strong>gthe night whereas Lobelia even at moderatetemperatures (≈16 o C) under natural conditionsexperiences anoxia <strong>in</strong> the roots (Møller & S<strong>and</strong>-Jensen 2011a). Furthermore, higherphotosynthesis of Littorella than Lobelia couldpromote higher organic carbon supplies to thesymbionts.AcknowledgementsWe thank Anubias (France) <strong>and</strong> Jan OlePedersen for provid<strong>in</strong>g non-mycorrhizalLittorella uniflora <strong>and</strong> Helene Rasmussen forprocess<strong>in</strong>g <strong>and</strong> count<strong>in</strong>g hyphal lengths <strong>in</strong><strong>sediment</strong> samples. We thank The Willum KannFoundation for f<strong>in</strong>ancial support to this studythrough The Centre of Excellence for Researchon Lake Restoration (CLEAR).References82
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