lm7h@bio.7u.nl).Nutrient resorption is an important process during leaf senescence, which helps plants to minimize nutrient losses. To quantify nutrient resorption, the parameter resorption efficiency is commonly used. This parameter describes the percentage of the nutrient pool withdrawn before leaf abscission. The nutrient pool is generally expressed on the basis of leaf mass or leaf area, assuming that these bases do not change during senescence. In this paper we firstly present a mathematical formula describing the effect of change in measurement basis on the difference between the real resorption efficiency (RRE) value and the measured resorption efficiency (MRE). This formula shows that even moderate senescence-related changes in a measurement basis can lead to considerable underestimation of RRE. Secondly, to estimate the general change in measurement basis we quantified leaf mass loss and leaf shrinkage during senescence from literature data. These data shows that mass loss percentages can be as high as 40%, and leaf shrinkage can be up to 20%. This level of change in basis seriously compromises the MRE when not corrected for. Using our formula and the reported average literature values of changes in leaf mass (21%) and leaf shrinkage (11%) during senescence, we calculated that the average RRE for nitrogen and phosphorous of terrestrial plants is 6% (leaf area) to 10% (leaf mass) higher than the 50%, respectively 52% as reported by Aerts (1996). This implies that nutrient resorption from senescing leaves is even more important for nutrient retention in terrestrial plants than thought so far. We advocate that preselecting leaves and monitoring the measurement basis throughout the duration of the experiment should minimize the difference between MRE and RRE.
Summary1 Plant growth at high-latitude sites is usually strongly nutrient-limited. The increased nutrient availability predicted in response to global warming may affect internal plant nutrient cycling, including nutrient resorption from senescing leaves. ) on nitrogen and phosphorus resorption efficiency and proficiency in six sub-arctic bog species, belonging to four different growth-forms, was studied in northern Sweden. 3 We hypothesized that while increased N supply would not affect N or P resorption efficiency, it would lead to lower N resorption proficiency (higher N concentrations in leaf litter) and higher P resorption proficiency (lower P concentrations in leaf litter). We also investigated whether the basis on which resorption was expressed (leaf mass, leaf area or unit leaf) influenced the patterns observed. 4 Contrasting with our hypothesis, a general trend of decreased N resorption efficiency occurred in response to increased N supply, but the expected decrease in N resorption proficiency was seen in all species except Betula nana . 5 P resorption efficiency did not change in four species ( B. nana , Empetrum hermaphroditum , Eriophorum vaginatum and Rubus chamaemorus ) but it decreased in Andromeda polifolia , and increased in Vaccinium uliginosum . P resorption proficiency showed the expected increase in only two species ( B. nana and V. uliginosum ). 6 Apart from P resorption efficiency, the different calculation methods generally produced similar responses of resorption efficiency and proficiency to N supply. 7 Increased N supply at high-latitude sites clearly leads to more N being returned to the soil through leaf litter production. However, decomposition of such litter will probably become P-limited. 8 Considerable interspecific differences in nutrient resorption proficiency were found, indicating that long-term changes in vegetation composition need to be considered when evaluating plant-mediated effects on ecosystem nutrient cycling in response to increased nutrient supply.
The relationship between biogeochemical process rates and microbial functional activity was investigated by analysis of the transcriptional dynamics of the key functional genes for methanogenesis (methyl coenzyme M reductase; mcrA) and methane oxidation (particulate methane monooxygenase; pmoA) and in situ methane flux at two peat soil field sites with contrasting net methane-emitting and -oxidizing characteristics. qPCR was used to quantify the abundances of mcrA and pmoA genes and transcripts at two soil depths. Total methanogen and methanotroph transcriptional dynamics, calculated from mcrA and pmoA gene : transcript abundance ratios, were similar at both sites and depths. However, a linear relationship was demonstrated between surface mcrA and pmoA transcript dynamics and surface flux rates at the methane-emitting and methane-oxidizing sites, respectively. Results indicate that methanotroph activity was at least partially substrate-limited at the methane-emitting site and by other factors at the methane-oxidizing site. Soil depth also contributed to the control of surface methane fluxes, but to a lesser extent. Small differences in the soil water content may have contributed to differences in methanogen and methanotroph activities. This study therefore provides a first insight into the regulation of in situ, field-level surface CH(4) flux at the molecular level by an accurate reflection of gene : transcript abundance ratios for the key genes in methane generation and consumption.
The efficacy and feasibility of annual harvesting of Phragmites australis and Typha latifolia shoots in autumn for nutrient removal was evaluated in a wetland system used for polishing sewage treatment plant (STP) effluent. Aboveground biomass and nutrient dynamics nutrient removal through harvest were studied in parallel ditches with stands of Phragmites or Typha that were mown in October during two successive years. The inflow rate of STP effluent to the ditches was experimentally varied, resulting in pairs of ditches with mean hydraulic retention times (HRT) of 0.3, 0.8, 2.3, and 9.3 days, corresponding to N and P mass loading rates of 122-4190 g N m(-2) yr(-1) and 28.3-994 g P m(-2) yr(-1). Nitrogen and P removal efficiency by harvest of Phragmites and Typha shoots in October increased with increasing HRT, despite the opposite HRT effect on N and P standing stocks. This removal through harvest appeared to be useful in treatment wetlands with N and P mass loading rates lower than approximately 120 g N m(-2) yr(-1) and 30 g P m(-2) yr(-1), corresponding to a HRT of roughly 9 days in the ditches of this wetland system. At the HRT of 9.3 days, the annual mass input to the ditches was reduced through the harvest by 7.0-11% and 4.5 -9.2% for N and P, respectively. At the higher nutrient mass loading rates, the nutrient removal through harvest was insignificant compared to the mass inputs. The vitality of Phragmites and Typha, measured as maximum aboveground biomass, was not affected by the annual cutting of the shoots in autumn over two years. The Typha stands yielded higher N and P removal efficiencies through shoot harvest than the Phragmites stands, which was largely the result of lower decreases in N and P standing stocks between August and October. This difference in nutrient standing stocks between the two species was caused by a combined effect of greater decreases in nutrient concentrations largely due to higher nutrient retranslocation efficiencies of Phragmites plants and greater reductions in shoot Phragmites biomass because of leaf fall and mass resorption. Nutrient removal by harvesting Phragmites shoots can probably be doubled without a reduction in vitality of the stands by advancing the harvest date to mid-September, which would at least approach the nutrient removal by harvesting Typha shoots in October. Phragmites also may be more profitable in very low-loaded wetland systems because the vigor of Typha stands seemed to be more sensitive to a lower nutrient availability at N and P mass input rates lower than the range indicated.
We studied the effects of elevated CO 2 (180 -200 ppmv above ambient) on growth and chemistry of three moss species (Sphagnum palustre, S. recurvum and Polytrichum commune) in a lowland peatland in the Netherlands. Thereto, we conducted both a greenhouse experiment with both Sphagnum species and a field experiment with all three species using MiniFACE (Free Air CO 2 Enrichment) technology during 3 years. The greenhouse experiment showed that Sphagnum growth was stimulated by elevated CO 2 in the short term, but that in the longer term ( ‡1 year) growth was probably inhibited by low water tables and/or downregulation of photosynthesis. In the field experiment, we did not find significant changes in moss abundance in response to elevated CO 2 , although CO 2 enrichment appeared to reduce S. recurvum abundance. Both Sphagnum species showed stronger responses to spatial variation in hydrology than to increased atmospheric CO 2 concentrations. Polytrichum was insensitive to changes in hydrology. Apart from the confounding effects of hydrology, the relative lack of growth response of the moss species may also have been due to the relatively small increase in assimilated CO 2 as achieved by the experimentally added CO 2 . We calculated that the added CO 2 contributed at most 32% to the carbon assimilation of the mosses, while our estimates based on stable C isotope data even suggest lower contributions for Sphagnum (24 -27%). Chemical analyses of the mosses showed only small elevated CO 2 effects on living tissue N concentration and C/N ratio of the mosses, but the C/N ratio of Polytrichum was substantially lower than those of the Sphagnum species. Continuing expansion of Polytrichum at the expense of Sphagnum could reduce the C sink function of this lowland Sphagnum peatland, and similar ones elsewhere, as litter decomposition rates would probably be enhanced. Such a reduction in sink function would be driven mostly by increased atmospheric N deposition, water table regulation for agricultural purposes and land management to preserve the early successional stage (mowing, tree and shrub removal), since these anthropogenic factors will probably exert a greater control on competition between Polytrichum and Sphagnum than increased atmospheric CO 2 concentrations.
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