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Climate warming poses major threats to temperate forests, but the response of tree root metabolism has largely remained unclear. We examined the impact of long‐term soil warming (>14 years, +4°C) on the fine root metabolome across three seasons for 2 years in an old spruce forest, using a liquid chromatography‐mass spectrometry platform for primary metabolite analysis. A total of 44 primary metabolites were identified in roots (19 amino acids, 12 organic acids and 13 sugars). Warming increased the concentration of total amino acids and of total sugars by 15% and 21%, respectively, but not organic acids. We found that soil warming and sampling date, along with their interaction, directly influenced the primary metabolite profiles. Specifically, in warming plots, concentrations of arginine, glycine, lysine, threonine, tryptophan, mannose, ribose, fructose, glucose and oxaloacetic acid increased by 51.4%, 19.9%, 21.5%, 19.3%, 22.1%, 23.0%, 38.0%, 40.7%, 19.8% and 16.7%, respectively. Rather than being driven by single compounds, changes in metabolite profiles reflected a general up‐ or downregulation of most metabolic pathway network. This emphasises the importance of metabolomics approaches in investigating root metabolic pathways and understanding the effects of climate change on tree root metabolism. Summary statement: Soil warming increases the content of amino acids and sugars in mature Norway spruce (Picea abies) fine roots, but not of organic acids. Moreover, we found that the changes in metabolite profiles were not driven by individual compounds but by a general up‐ or downregulation of most primary metabolites. [ABSTRACT FROM AUTHOR]

Microbial necromass carbon (MNC) accounts for a large fraction of soil organic carbon (SOC) in terrestrial ecosystems. Yet our understanding of the fate of this large carbon pool under long‐term warming is uncertain. Here, we show that 14 years of soil warming (+4°C) in a temperate forest resulted in a reduction in MNC by 11% (0–10 cm) and 33% (10–20 cm). Warming caused a decrease in the content of MNC due to a decline in microbial biomass carbon and reduced microbial carbon use efficiency. This reduction was primarily caused by warming‐induced limitations in available soil phosphorus, which, in turn, constrained the production of microbial biomass. Conversely, warming increased the activity of soil extracellular enzymes, specifically N‐acetylglucosaminidase and leucine aminopeptidase, which accelerated the decomposition of MNC. These findings collectively demonstrate that decoupling of MNC formation and decomposition underlie the observed MNC loss under climate warming, which could affect SOC content in temperate forest ecosystems more widespread. [ABSTRACT FROM AUTHOR]

The major driving factors of soil CO2 production - substrate supply, temperature, and water content - vary vertically within the soil profile, with the greatest temporal variations of these factors usually near the soil surface. Several studies have demonstrated that wetting and drying of the organic horizon contributes to temporal variation in summertime soil CO2 efflux in forests, but this contribution is difficult to quantify. The objectives of this study were to partition CO2 production vertically in a mixed hardwood stand of the Harvard Forest, Massachusetts, USA, and then to use that partitioning to evaluate how the relative contributions of CO2 production by genetic soil horizon vary seasonally and interannually. We measured surface CO2 efflux and vertical soil profiles of CO2 concentration, temperature, water content, and soil physical characteristics. These data were applied to a model of effective diffusivity to estimate CO2 flux at the top of each genetic soil horizon and the production within each horizon. A sensitivity analysis revealed sources of uncertainty when applying a diffusivity model to a rocky soil with large spatial heterogeneity, especially estimates of bulk density and volumetric water content and matching measurements of profiles and surface fluxes. We conservatively estimate that the O horizon contributed 40-48% of the total annual soil CO2 efflux. Although the temperature sensitivity of CO2 production varied across soil horizons, the partitioning of CO2 production by horizon did not improve the overall prediction of surface CO2 effluxes based on temperature functions. However, vertical partitioning revealed that water content covaried with CO2 production only in the O horizon. Large interannual variations in estimates of O horizon CO2 production indicate that this layer could be an important transient interannual source or sink of ecosystem C. © 2006 Blackwell Publishing Ltd.

Soil moisture affects microbial decay of SOM and rhizosphere respiration (RR) in temperate forest soils, but isolating the response of soil respiration (SR) to summer drought and subsequent wetting is difficult because moisture changes are often confounded with temperature variation. We distinguished between temperature and moisture effects by simulation of prolonged soil droughts in a mixed deciduous forest at the Harvard Forest, Massachusetts. Roofs constructed over triplicate 5×5m2 plots excluded throughfall water during the summers of 2001 (168mm) and 2002 (344 mm), while adjacent control plots received ambient throughfall and the same natural temperature regime. In 2003, throughfall was not excluded to assess the response of SR under natural weather conditions after two prolonged summer droughts. Throughfall exclusion significantly decreased mean SR rate by 53mgCm-2h-1 over 84 days in 2001, and by 68mgCm-2h-1 over 126 days in 2002, representing 10-30% of annual SR in this forest and 35-75% of annual net ecosystem exchange (NEE) of C. The differences in SR were best explained by differences in gravimetric water content in the Oi horizon (r250.69) and the Oe/Oa horizon (r250.60). Volumetric water content of the A horizon was not significantly affected by throughfall exclusion. The radiocarbon signature of soil CO2 efflux and of CO2 respired during incubations of O horizon, A horizon and living roots allowed partitioning of SR into contributions from young C substrate (including RR) and from decomposition of older SOM. RR (root respiration and microbial respiration of young substrates in the rhizosphere) made up 43-71% of the total C respired in the control plots and 41-80% in the exclusion plots, and tended to increase with drought. An exception to this trend was an interesting increase in CO2 efflux of radiocarbon-rich substrates during a period of abundant growth of mushrooms. Our results suggest that prolonged summer droughts decrease primarily heterotrophic respiration in the O horizon, which could cause increases in the storage of soil organic carbon in this forest. However, the C stored during two summers of simulated drought was only partly released as increased respiration during the following summer of natural throughfall. We do not know if this soil C sink during drought is transient or long lasting. In any case, differential decomposition of the O horizon caused by interannual variation of precipitation probably contributes significantly to observed interannual variation of NEE in temperate forests. © 2005 Blackwell Publishing Ltd.

Many moss species are associated with nitrogen (N)‐fixing bacteria (diazotrophs) that support the N supply of mosses. Our knowledge relates primarily to pristine ecosystems with low atmospheric N input, but knowledge of biological N fixation (BNF) and diazotrophic communities in mosses in temperate forests with high N deposition is limited. We measured BNF rates using the direct stable isotope method and studied the total and potentially active diazotrophic communities in two abundant mosses, Brachythecium rutabulum and Hypnum cupressiforme, both growing on lying deadwood trunks in 25 temperate forest sites. BNF rates in both mosses were similar to those observed in moss species of pristine ecosystems. H. cupressiforme fixed three times more N2 and exhibited lower diazotrophic richness than B. rutabulum. Frankia was the most prominent diazotroph followed by cyanobacteria Nostoc. Manganese, iron, and molybdenum contents in mosses were positively correlated with BNF and diazotrophic communities. Frankia maintained high BNF rates in H. cupressiforme and B. rutabulum even under high chronic N deposition in Central European forests. Moss N concentration and 15N abundance indicate a rather minor contribution of BNF to the N nutrition of these mosses. [ABSTRACT FROM AUTHOR]