Peat bogs have historically represented exceptional carbon (C) sinks because of their extremely low decomposition rates and consequent accumulation of plant remnants as peat. Among the factors favoring that peat accumulation, a major role is played by the chemical quality of plant litter itself, which is poor in nutrients and characterized by polyphenols with a strong inhibitory effect on microbial breakdown. Because bogs receive their nutrient supply solely from atmospheric deposition, the global increase of atmospheric nitrogen (N) inputs as a consequence of human activities could potentially alter the litter chemistry with important, but still unknown, effects on their C balance. Here we present data showing the decomposition rates of recently formed litter peat samples collected in nine European countries under a natural gradient of atmospheric N deposition from Ϸ0.2 to 2 g⅐m ؊2 ⅐yr ؊1 . We found that enhanced decomposition rates for material accumulated under higher atmospheric N supplies resulted in higher carbon dioxide (CO2) emissions and dissolved organic carbon release. The increased N availability favored microbial decomposition (i) by removing N constraints on microbial metabolism and (ii) through a chemical amelioration of litter peat quality with a positive feedback on microbial enzymatic activity. Although some uncertainty remains about whether decay-resistant Sphagnum will continue to dominate litter peat, our data indicate that, even without such changes, increased N deposition poses a serious risk to our valuable peatland C sinks.decomposition ͉ global change ͉ litter peat ͉ CO2 P eatlands cover 2-3% of the land's surface, store approximately one-third of all soil carbon (C) (390-455 Pg), and currently act as sinks for atmospheric C (1, 2). The ability of peatlands to sequester atmospheric C resides in the long-term accumulation of partially decomposed organic matter (i.e., peat). Indeed, acidic water conditions, low soil temperature, frequent waterlogging, and low nutrient quality of plant litter impair decomposition of plant litter, favoring its accumulation (3). In peatlands exclusively fed by atmospheric deposition (i.e., bogs) (1), the accumulated peat is dominated by the remnants of the mosses of the genus Sphagnum, which produce a litter poor in nutrients and highly enriched in organochemical compounds such as uronic acids (4) and polyphenols (5) with a strong inhibitory effect on microbial activity and vascular plants (3). As such, Sphagnum plants form the bulk of living and dead biomass in bog ecosystems (3).Because of the strict dependence of bogs on atmospheric deposition as a source of nutrients (1), the increasing availability of biologically reactive nitrogen (N) from industrial and agricultural activities (6) could potentially alter the chemical quality of plant litter with consequent effects on the amount of C released during litter decomposition. Accordingly, an understanding of the mechanisms of bog soil C response to changing N availability is essential for assessing the capa...

Article (refereed) -postprintTipping, E.; Benham, S.; Boyle, J.F.; Crow, P.; Davies, J.; Fischer, U.; Guyatt, H.; Helliwell, R.; Jackson-Blake, L.; Lawlor, A.J.; Monteith, D.T.; Rowe, E.C.; Toberman, H. 2014. Atmospheric deposition of phosphorus to land and freshwater. Environmental Science: Processes and Impacts, 16 (7). 1608-1617. 10.1039/c3em00641g Contact CEH NORA team at noraceh@ceh.ac.ukThe NERC and CEH trademarks and logos ('the Trademarks') are registered trademarks of NERC in the UK and other countries, and may not be used without the prior written consent of the Trademark owner. Oceania, and South-Central America. The deposition rates are log-normally distributed, 30 and for the whole data set the geometric mean deposition rates are 0.027, 0.019 and 31 0.14 g m -2 a -1 for TP, FTP and PO 4 -P respectively. At smaller scales there is little 32 systematic spatial variation, except for high deposition rates at some sites in Germany, 33 likely due to local agricultural sources. In cases for which PO 4 -P was determined as well 34 as one of the other forms of P, strong parallels between logarithmic values were found. 35Based on the directly-measured deposition rates to land, and published estimates of P 36 deposition to the oceans, we estimate a total annual transfer of P to and from the 37 atmosphere of 3.7 Tg. However, much of the phosphorus in larger particles (principally 38 primary biological aerosol particles) is probably redeposited near to its origin, so that 39 long-range transport, important for tropical forests, large areas of peatland and the 40 oceans, mainly involves fine dust from deserts and soils, as described by the simulations 41of Mahowald et al. (Global Biogeochemical Cycles 22, GB4026, 2008). We suggest that 42 local release to the atmosphere and subsequent deposition bring about a pseudo-43 diffusive redistribution of P in the landscape, with P-poor ecosystems, for example 44 ombrotrophic peatlands and oligotrophic lakes, gaining at the expense of P-rich ones. 45Simple calculations suggest that atmospheric transport could bring about significant local 46 redistribution of P among terrestrial ecosystems.Although most atmospherically 47 transported P is natural in origin, local transfers from fertilised farmland to P-poor 48 ecosystems may be significant, and this requires further research. 49 50

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Abstract. Peatlands are carbon (C) storage ecosystems sustained by a high water table (WT). High WT creates anoxic conditions that suppress the activity of aerobic decomposers and provide conditions for peat accumulation. Peatland function can be dramatically affected by WT drawdown caused by climate and/or land-use change. Aerobic decomposers are directly affected by WT drawdown through environmental factors such as increased oxygenation and nutrient availability. Additionally, they are indirectly affected via changes in plant community composition and litter quality. We studied the relative importance of direct and indirect effects of WT drawdown on aerobic decomposer activity in plant litter at two stages of decomposition (incubated in the field for 1 or 2 years). We did this by profiling 11 extracellular enzymes involved in the mineralization of organic C, nitrogen (N), phosphorus (P) and sulphur. Our study sites represented a three-stage chronosequence from pristine to shortterm (years) and long-term (decades) WT drawdown conditions under two nutrient regimes (bog and fen). The litter types included reflected the prevalent vegetation: Sphagnum mosses, graminoids, shrubs and trees.Litter type was the main factor shaping microbial activity patterns and explained about 30 % of the variation in enzyme activities and activity allocation. Overall, enzyme activities were higher in vascular plant litters compared to Sphagnum litters, and the allocation of enzyme activities towards C or nutrient acquisition was related to the initial litter quality Correspondence to: P. Straková (petra.strakova@helsinki.fi) (chemical composition). Direct effects of WT regime, site nutrient regime and litter decomposition stage (length of incubation period) summed to only about 40 % of the litter type effect. WT regime alone explained about 5 % of the variation in enzyme activities and activity allocation. Generally, enzyme activity increased following the long-term WT drawdown and the activity allocation turned from P and N acquisition towards C acquisition. This caused an increase in the rate of litter decomposition. The effects of the shortterm WT drawdown were minor compared to those of the long-term WT drawdown: e.g., the increase in the activity of C-acquiring enzymes was up to 120 % (bog) or 320 % (fen) higher after the long-term WT drawdown compared to the short-term WT drawdown. In general, the patterns of microbial activity as well as their responses to WT drawdown depended on peatland type: e.g., the shift in activity allocation to C-acquisition was up to 100 % stronger at the fen compared to the bog.Our results imply that changes in plant community composition in response to persistent WT drawdown will strongly affect the C dynamics of peatlands. The predictions of decomposer activity under changing climate and/or land-use thus cannot be based on the direct effects of the changed environment only, but need to consider the indirect effects of environmental changes: the changes in plant community composition, their dependen...

Fire is a major driver of ecosystem change and can disproportionately affect the cycling of different nutrients. Thus, a stoichiometric approach to investigate the relationships between nutrient availability and microbial resource use during decomposition is likely to provide insight into the effects of fire on ecosystem functioning. We conducted a field litter bag experiment to investigate the long-term impact of repeated fire on the stoichiometry of leaf litter C, N and P pools, and nutrient-acquiring enzyme activities during decomposition in a wet sclerophyll eucalypt forest in Queensland, Australia. Fire frequency treatments have been maintained since 1972, including burning every 2 years (2yrB), burning every 4 years (4yrB) and no burning (NB). C : N ratios in freshly fallen litter were 29-42% higher and C : P ratios were 6-25% lower for 2yrB than NB during decomposition, with correspondingly lower 2yrB N : P ratios (27-32) than for NB (34-49). Trends in litter soluble and microbial N : P ratios were similar to the overall litter N : P ratios across fire treatments. Consistent with these, the ratio of activities for N-acquiring to P-acquiring enzymes in litter was higher for 2yrB than NB, whereas 4yrB was generally intermediate between 2yrB and NB. Decomposition rates of freshly fallen litter were significantly lower for 2yrB (72 AE 2% mass remaining at the end of experiment) than for 4yrB (59 AE 3%) and NB (62 AE 3%), a difference that may be related to effects of N limitation, lower moisture content, and/or litter C quality. Results for older mixed-age litter were similar to those for freshly fallen litter although treatment differences were less pronounced. Overall, these findings show that frequent fire (2yrB) decoupled N and P cycling, as manifested in litter C : N : P stoichiometry and in microbial biomass N : P ratio and enzymatic activities. Furthermore, these data indicate that fire induced a transient shift to N-limited ecosystem conditions during the postfire recovery phase.

Natural moisture limitation during summer drought can constitute a stress for microbial communities in soil. Given globally predicted increases in drought frequency, there is an urgent need for a greater understanding of the effects of drought events on soil microbial processes. Using a long-term field-scale drought manipulation experiment at Clocaenog, Wales, UK, we analysed fungal community dynamics, using internal transcribed spacer-denaturing gradient gel electrophoresis (DGGE), over a 1-year period in the 6th year of drought manipulation. Ambient seasonality was found to be the dominant factor driving variation in fungal community dynamics. The summer drought manipulation resulted in a significant decline in the abundance of dominant fungal species, both independently of, and in interaction with, this seasonal variation. Furthermore, soil moisture was significantly correlated with the changes in fungal diversity over the drought manipulation period. While the relationship between species diversity and functional diversity remains equivocal, phenol oxidase activity was decreased by the summer drought conditions and there was a significant correlation with the decline of DGGE band richness among the most dominant fungal species during the drought season. Climatically driven events such as droughts may have significant implications for fungal community diversity and therefore, have the potential to interfere with crucial ecosystem processes, such as organic matter decomposition.

The effects of 4 years of simulated nitrogen deposition, as nitrate (NO 3 À ) and ammonium (NH 4 1 ), on microbial carbon turnover were studied in an ombrotrophic peatland. We investigated the mineralization of simple forms of carbon using MicroRespt measurements (a multiple substrate induced respiration technique) and the activities of four soil enzymes involved in the decomposition of more complex forms of carbon or in nutrient acquisition: Nacetyl-glucosaminidase (NAG), cellobiohydrolase (CBH), acid phosphatase (AP), and phenol oxidase (PO). The potential mineralization of labile forms of carbon was significantly enhanced at the higher N additions, especially with NH 4 1 amendments, while potential enzyme activities involved in breakdown of more complex forms of carbon or nutrient acquisition decreased slightly (NAG and CBH) or remained unchanged (AP and PO) with N amendments. This study also showed the importance of distinguishing between NO 3 À and NH 4 1 amendments, as their impact often differed. It is possible that the limited response on potential extracellular enzyme activity is due to other factors, such as limited exposure to the added N in the deeper soil or continued suboptimal functioning of the enzymes due to the low pH, possibly via the inhibitory effect of low phenol oxidase activity.

Drainage for forestry has been amongst the most extensive of land management practices applied to northern latitude peatlands, particularly in northern Europe. Extracellular phenol oxidases play an important role in the carbon cycle of soils. This study investigated the effects of long-term (45 years) drainage for forestry upon surface peat extracellular phenol oxidase activity, soluble phenolic concentrations and pH at ombrotrophic bog, oligotrophic fen and mesotrophic fen sites at a Finnish mire complex. Phenol oxidase activity was reduced by drainage at all three sites. Phenol oxidase activity was positively correlated with peat pH across all sites irrespective of drainage treatment, suggesting that pH is a major factor influencing peat phenol oxidase activity at the mire complex. Peat pH became more acidic with drainage at the fen sites, and it is likely that this contributed to the suppression of peat phenol oxidase activity. The reduction of peat phenol oxidase activity with drainage was accompanied by increased concentrations of water-soluble phenolics at all three sites, and the potential contribution of this to changes in peat carbon stocks following drainage is discussed.