Organic matter inputs shift soil enzyme activity and allocation patterns in a wet tropical forest

Soil extracellular enzymes mediate organic matter turnover and nutrient cycling yet remain little studied in one of Earth’s most rapidly changing, productive biomes: tropical forests. Using a long-term leaf litter and throughfall manipulation, we explored relationships between organic matter (OM) inputs, soil chemical properties and enzyme activities in a lowland tropical forest. We assayed six hydrolytic soil enzymes responsible for liberating carbon (C), nitrogen (N) and phosphorus (P), calculated enzyme activities and ratios in control plots versus treatments, and related these to soil biogeochemical variables. While leaf litter addition and removal tended to increase and decrease enzyme activities per gram soil, respectively, shifts in enzyme allocation patterns implied changes in relative nutrient constraints with altered OM inputs. Enzyme activity ratios in control plots suggested strong belowground P constraints; this was exacerbated when litter inputs were curtailed. Conversely, with double litter inputs, increased enzymatic investment in N acquisition indicated elevated N demand. Across all treatments, total soil C correlated more strongly with enzyme activities than soluble C fluxes, and enzyme ratios were sensitive to resource stoichiometry (soil C:N) and N availability (net N mineralization). Despite high annual precipitation in this site (MAP ~5 m), soil moisture positively correlated with five of six enzymes. Our results suggest resource availability regulates tropical soil enzyme activities, soil moisture plays an additional role even in very wet forests, and relative investment in C, N and P degrading enzymes in tropical soils will often be distinct from higher latitude ecosystems yet is sensitive to OM inputs.     

Experimental removal and addition of leaf litter inputs reduces nitrate production and loss in a lowland tropical forest

Environmental perturbations such as changes in land use, climate, and atmospheric carbon dioxide concentrations may alter organic matter inputs to surface soils. While the carbon (C) cycle response to such perturbations has received considerable attention, potential responses of the soil nitrogen (N) cycle to changing organic matter inputs have been less well characterized. Changing litter inputs to surface to soils may alter the soil N cycle directly, by controlling N substrate availability, or indirectly, via interactions with soil C biogeochemistry. We investigated soil N-cycling responses to a leaf litter manipulation in a lowland tropical forest using isotopic and molecular techniques. Both removing and doubling leaf litter inputs decreased the size of the soil nitrate pool, gross nitrification rates, and the relative abundance of ammonia-oxidizing microorganisms. Gross nitrification rates were correlated with the relative abundance of ammonia-oxidizing archaea, and shifts in the N-cycling microbial community composition correlated with concurrent changes in edaphic properties, notably pH and C:N ratios. These results highlight the importance of understanding coupled biogeochemical cycles in global change scenarios and suggest that environmental perturbations that alter organic matter inputs in tropical forests could reduce inorganic N losses to surface waters and the atmosphere by limiting nitrate production.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Endogenous versus exogenous nutrient affects C, N, and P dynamics in decomposing litters in mid-subtropical forests of China

The effect of nutrient availability on litter decomposition has been a major focus of global change ecology. The relative impacts of endogenous (litter) and exogenous (soil) nutrient availability remain unclear. We studied the nutrient dynamics of decomposition in litter from two species with contrasting litter nutrient contents and stoichiometry: Pinus massoniana and Castanopsis sclerophylla. During a 540-day field incubation, we manipulated exogenous nutrient levels by adding microbially available C (+C), N (+N), P (+P), and all three (+CNP) at 90-day intervals. Relative to the no-nutrient control (CK), nutrient additions decreased organic C retention in C. sclerophylla, with the greatest effect observed in +CNP. Nitrogen content in P. massoniana litter similarly increased with nutrient addition, particularly +P and +CNP. The P addition treatments also increased P content in the litter of both species. Nitrogen content in C. sclerophylla and organic C content in P. massoniana were unaffected by nutrient additions. The C/N and C/P ratios in decomposing C. sclerophylla litter were significantly lower in the CK treatment, while those of P. massoniana litter were influenced by the interaction of nutrient addition and decomposition time. Increased availability of C, N, and P individually and collectively alters nutrient release dynamics in decomposing foliar litter. Litter quality, as determined by source species, is a key determinant of the impact of exogenous nutrient inputs. A stronger effect of P addition than N addition indicates a relatively N-rich and P-poor ecosystem.     

Indirect Effects of Nitrogen Amendments on Organic Substrate Quality Increase Enzymatic Activity Driving Decomposition in a Mesic Grassland

The fate of soil organic carbon (SOC) is determined, in part, by complex interactions between the quality of plant litter inputs, nutrient availability, and the microbial communities that control decomposition rates. This study explores these interactions in a mesic grassland where C and nitrogen (N) availability and plant litter quality have been manipulated using both fertilization and haying for 7 years. We measured a suite of soil parameters including inorganic N, extractable organic C and N (EOC and EON), soil moisture, extracellular enzyme activity (EEA), and the isotopic composition of C and N in the microbial biomass and substrate sources. We use these data to determine how the activity of microbial decomposers was influenced by varying levels of substrate C and N quality and quantity and to explore potential mechanisms explaining the fate of enhanced plant biomass inputs with fertilization. Oxidative EEA targeting relatively recalcitrant C pools was not affected by fertilization. EEA linked to the breakdown of relatively labile C rich substrates exhibited no relationship with inorganic N availability but was significantly greater with fertilization and associated increases in substrate quality. These increases in EEA were not related to an increase in microbial biomass C. The ratio of hydrolytic C:N acquisition enzymes and δ¹³C and δ¹⁵N values of microbial biomass relative to bulk soil C and N, or EOC and EON suggest that microbial communities in fertilized plots were relatively C limited, a feature likely driving enhanced microbial efforts to acquire C from labile sources. These data suggest that in mesic grasslands, enhancements in biomass inputs and quality with fertilization can prompt an increase in EEA within the mineral soil profile with no significant increases in microbial biomass. Our work helps elucidate the microbially mediated fate of enhanced biomass inputs that are greater in magnitude than the associated increases in mineral soil organic matter.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

Nitrogen Translocation to Fresh Litter in Northern Hardwood Forest

Nitrogen immobilization in fresh litter represents a significant N flux in forest ecosystems, and changes in this process resulting from atmospheric N deposition could have important implications for ecosystem responses. We conducted two leaf decay experiments, using ¹⁵N-labeled sugar maple leaf litter, to quantify N transport from old litter and soil to fresh litter during early stages of decomposition, and we examined the influence of litter N concentration and soil N availability on upward N transfer in a northern hardwood forest. After one year of decay, the average N transfer from soil to fresh litter (2.63 mg N g^?1 litter) was much higher than the N transfer from older litter (1- to 2-years-old) to fresh litter (0.37 mg N g^?1 litter). We calculated the ratio of annual N transfer per unit of excess ¹⁵N pool for these two N sources. The ratio was not significantly different between old litter and soil, suggesting that fungi utilize N in the old litter and mineral soil pools for transport to decaying fresh litter with similar efficiency. Initial litter N concentration had a significant effect on upward N flux into decaying leaf litter, whereas no effect of soil N fertilization was observed. Reduction in the flux from soil to fresh litter owing to anthropogenic N inputs probably contributes significantly to changing soil N dynamics. Future work is needed on fungal N acquisition and transport as well as the fungal taxa involved in this process and their responses to changing environments.     

Global Positive Effects of Litter Inputs on Soil Nitrogen Pools and Fluxes

Ecosystems - Soil nitrogen (N) pools and fluxes are strongly influenced by litter inputs in natural ecosystems. However, general responses of soil N pools and fluxes to litter inputs from both...     

Identification of General Patterns of Nutrient and Labile Carbon Control on Soil Carbon Dynamics Across a Successional Gradient

Carbon (C) inputs and nutrient availability are known to affect soil organic carbon (SOC) stocks. However, general rules regarding the operation of these factors across a range of soil nutrient availabilities and substrate qualities are unidentified. “Priming” (stimulated decomposition by labile C inputs) and ‘preferential substrate utilization’ (retarded decomposition due to shifts in community composition towards microbes that do not mineralize SOC) are two hypotheses to explain effects of labile C additions on SOC dynamics. For effects of nutrient additions (nitrogen and phosphorus) on SOC dynamics, the stoichiometric (faster decomposition of materials of low carbon-to-nutrient ratios) and ‘microbial mining’ (that is, reduced breakdown of recalcitrant C forms for nutrients under fertile conditions) hypotheses have been proposed. Using the natural gradient of soil nutrient availability and substrate quality of a chronosequence, combined with labile C and nutrient amendments, we explored the support for these contrasting hypotheses. Additions of labile C, nitrogen (N), phosphorus (P), and combinations of C and N and C and P were applied to three sites: 2-year fallow grassland, mature grassland and forest, and the effects of site and nutrient additions on litter decomposition and soil C dynamics were assessed. The response to C addition supported the preferential substrate hypothesis for easily degradable litter C and the priming hypothesis for SOC, but only in nitrogen-enriched soils of the forest site. Responses to N addition supported the microbial mining hypothesis irrespective of C substrate (litter or SOC), but only in the forest site. Further, P addition effects on SOC support the stoichiometric hypothesis; P availability appeared key to soil C release (priming) in the forest site if labile C and N is available. These results clearly link previously contrasting hypotheses of the factors controlling SOC with the natural gradient in litter quality and nutrient availability that exists in ecosystems at different successional stages. A holistic theory that incorporates this variability of responses, due to different mechanisms, depending on nutrient availability and substrate quality is essential for devising management strategies to safeguard soil C stocks.     

Nutrient Addition Prompts Rapid Destabilization of Organic Matter in an Arctic Tundra Ecosystem

Nutrient availability in the arctic is expected to increase in the next century due to accelerated decomposition associated with warming and, to a lesser extent, increased nitrogen deposition. To explore how changes in nutrient availability affect ecosystem carbon (C) cycling, we used radiocarbon to quantify changes in belowground C dynamics associated with long-term fertilization of graminoid-dominated tussock tundra at Toolik Lake, Alaska. Since 1981, yearly fertilization with nitrogen (N) and phosphorus (P) has resulted in a shift to shrub-dominated vegetation. These combined changes have altered the quantity and quality of litter inputs, the vertical distribution and dynamics of fine roots, and the decomposition rate of soil organic C. The loss of C from the deep organic and mineral soil has more than offset the C accumulation in the litter and upper organic soil horizons. In the litter and upper organic horizons, radiocarbon measurements show that increased inputs resulted in overall C accumulation, despite being offset by increased decomposition in some soil pools. To reconcile radiocarbon observations in the deeper organic and mineral soil layers, where most of the ecosystem C loss occurred, both a decrease in input of new root material and a dramatic increase of decomposition rates in centuries-old soil C pools were required. Therefore, with future increases in nutrient availability, we may expect substantial losses of C which took centuries to accumulate.     

The Effect of Phosphorus Availability on Decomposition Dynamics in a Seasonal Lowland Amazonian Forest

Once the weathering of parent material ceases to supply significant inputs of phosphorus (P), vegetation depends largely on the decomposition of litter and soil organic matter and the associated mineralization of organic P forms to provide an adequate supply of this essential nutrient. At the same time, the decomposition of litter is often characterized by the immobilization of nutrients, suggesting that nutrient availability is a limiting factor for this process. Immobilization temporally decouples nutrient mineralization from decomposition and may play an important role in nutrient retention in low-nutrient ecosystems. In this study, we used a common substrate to study the effects of native soil P availability as well as artificially elevated P availability on litter decomposition rates in a lowland Amazonian rain forest on highly weathered soils. Although both available and total soil P pools varied almost three fold across treatments, there was no significant difference in decomposition rates among treatments. Decomposition was rapid in all treatments, with approximately 50% of the mass lost over the 11-month study period. Carbon (C) and nitrogen (N) remaining and C:N ratios were the most effective predictors of amount of mass remaining at each time point in all treatments. Fertilized treatments showed significant amounts of P immobilization (P < 0.001). By the final collection point, the remaining litter contained a quantity equivalent to two-thirds of the initial P and N, even though only half of the original mass remained. In these soils, immobilization of nutrients in the microbial biomass, late in the decomposition process, effectively prevents the loss of essential nutrients through leaching or occlusion in the mineral soil.     

Relationships among phosphorus, molybdenum and free-living nitrogen fixation in tropical rain forests: results from observational and experimental analyses

Biological nitrogen (N) fixation is the primary source of “new” N to unmanaged ecosystems, and recent analyses suggest that terrestrial N inputs via free-living N fixation may be more important than previously assumed. This may be particularly true in some tropical rain forests, where free-living fixation could outpace symbiotic N fixation to represent the dominant source of new N inputs. However, our understanding of the controls over free-living N fixation in tropical rain forests remains poor, which directly constrains our ability to predict how N cycling will respond to changing environmental conditions. Although both phosphorus (P) and molybdenum (Mo) availability have been shown to limit free-living N fixation rates in the tropics, few studies have simultaneously explored P versus Mo limitation or the potential importance of P × Mo interactions. Here, an archived set of foliar, litter, and soil samples from a Costa Rican tropical rain forest provided an opportunity to simultaneously assess the relative strength of P versus Mo relationships with free-living N fixation rates. We also conducted a short-term, full-factorial (P × Mo) litter incubation experiment to directly assess nutrient limitation, allowing us to explore P and Mo controls over free-living N fixation rates using both observational and experimental approaches. We previously showed that N fixation rates were positively correlated with P concentrations in all substrates and, using the archived samples, we now show that Mo concentrations correlated with N fixation only in canopy leaves (where total Mo concentrations were extremely low). Likewise, fertilization with P alone (and not Mo) stimulated leaf litter N fixation rates. Thus, our results suggest that P availability dominantly controls free-living N fixation at this site, and when taken with data from other studies, our results suggest that attempts to identify “the nutrient” that limits N fixation in “the tropics” may be misguided. Rather, nutrient controls over free-living N fixation appear to be more nuanced—and the true nature of nutrient limitation to N fixation likely varies over a variety of scales across the vast tropical rain forest biome.     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Contrasting Effects of Substrate and Fertilizer Nitrogen on the Early Stages of Litter Decomposition

Commonly observed positive correlations between litter nitrogen (N) concentrations and decomposition rates suggest that N frequently limits decomposition in its early stages. However, numerous studies have found little, if any, effect of N fertilization on decomposition. I directly compared internal substrate N and externally supplied inorganic N effects on decomposition in sites varying in soil N availability. I decomposed eight substrates (with initial %N from 0–2.5) in control and N-fertilized plots at eight grassland and forest sites in central Minnesota. N fertilization increased decomposition at only two of eight sites, even though decomposition was positively related to litter N at all sites and to soil N availability across sites. The effect of externally supplied N on decomposition was independent of litter N concentration, but was greater at sites with low N availability. The inconsistent effects of substrate and externally supplied N may have arisen because decomposers use organic N preferentially as an N source; because inorganic N availability across sites or with fertilization induced changes in microbial community attributes (for example, lower C:N or greater efficiency) that reduced the response of decomposition to increased inorganic N supply; or because the positive correlation between litter N or site N availability with decomposition was spurious, caused by tight correlations between litter or site N and some other factor that truly limited decomposition. These inconsistent effects of substrate N and external N supply on decomposition suggest that the oft-observed relationship between litter N and decomposition may not indicate N limitation of decomposition.     

Feedback from soil inorganic nitrogen on soil organic matter mineralisation and growth in a boreal forest ecosystem

Current nitrogen (N) deposition rates are considerably higher than during pre-industrial times and the growing interest in forest fertilisation requires better understanding of how the N and carbon (C) cycles interact. This study is based on experimental data showing how Scots pine (Pinus sylvestris L.) forests respond to single or consecutive pulse doses of N. The data were used to support the implementation of a dynamic feedback mechanism in the Q model, allowing for changes in soil N availability to regulate the rate of decomposer efficiency. Simulations of the long-term effects of slowly increasing N deposition with and without dynamic decomposer efficiency were then compared. Both versions of the model accurately predicted the response of tree growth to N fertilisation. Slowly increasing inputs of N over a century in the modified version acted on the inputs and outputs of soil C in opposing ways: (a) rate of litter input slowed down because more N was retained in the soil and thus not available for tree growth; (b) rate of C output, through soil heterotrophic respiration, was also gradually reduced due to increasing decomposer efficiency, although not enough to sufficiently balance the reduced litter input. Accurate prediction of the amount of added N retained in the ecosystem seems to be one of the key issues for estimating enhanced C sequestration.     

Effect of soil microorganisms and labile C availability on soil respiration in response to litter inputs in forest ecosystems: A meta‐analysis

Litter inputs can influence soil respiration directly through labile C availability and, indirectly, through the activity of soil microorganisms and modifications in soil microclimate; however, their relative contributions and the magnitude of any effect remain poorly understood. We synthesized 66 recently published papers on forest ecosystems using a meta‐analysis approach to investigate the effect of litter inputs on soil respiration and the underlying mechanisms involved. Our results showed that litter inputs had a strong positive impact on soil respiration, labile C availability, and the abundance of soil microorganisms, with less of an impact related to soil moisture and temperature. Overall, soil respiration was increased by 36% and 55%, respectively, in response to natural and doubled litter inputs. The increase in soil respiration induced by litter inputs showed a tendency for coniferous forests (50.7%)> broad‐leaved forests (41.3%)> mixed forests (31.9%). This stimulation effect also depended on stand age with 30‐ to 100‐year‐old forests (53.3%) and ≥100‐year‐old forests (50.2%) both 1.5 times larger than ≤30‐year‐old forests (34.5%). Soil microbial biomass carbon and soil dissolved organic carbon increased by 21.0%‐33.6% and 60.3%‐87.7%, respectively, in response to natural and doubled litter inputs, while soil respiration increased linearly with corresponding increases in soil microbial biomass carbon and soil dissolved organic carbon. Natural and doubled litter inputs increased the total phospholipid fatty acid (PLFA) content by 6.6% and 19.7%, respectively, but decreased the fungal/bacterial PLFA ratio by 26.9% and 18.7%, respectively. Soil respiration also increased linearly with increases in total PLFA and decreased linearly with decreases in the fungal/bacterial PLFA ratio. The contribution of litter inputs to an increase in soil respiration showed a trend of total PLFA > fungal/bacterial PLFA ratio > soil dissolved organic carbon > soil microbial biomass carbon. Therefore, in addition to forest type and stand age, labile C availability and soil microorganisms are also important factors that influence soil respiration in response to litter inputs, with soil microorganisms being more important than labile C availability.     

Anthropogenic N Deposition Increases Soil C Storage by Decreasing the Extent of Litter Decay: Analysis of Field Observations with an Ecosystem Model

Recent meta-analyses of experimental studies simulating increased anthropogenic nitrogen (N) deposition in forests reveal greater soil carbon (C) storage under elevated levels of atmospheric N deposition. However, these effects have not yet been included in ecosystem-scale models of soil C and N cycling and it is unclear whether increased soil C storage results from slower decomposition rates or a reduced extent of decomposition (for example, an increase in the amount of litter entering slowly decaying humus pools). To test these alternatives, we conducted a meta-analysis of litter decomposition data. We then used the results from our meta-analysis to model C and N cycling in four sugar maple forests in Michigan using an ecosystem process model (TRACE). We compared model results testing our alternative hypotheses to field data on soil C storage from a 17-year N deposition experiment. Using data from published litter decomposition studies in forests, we determined that, on average, exogenous N inputs decreased lignin decomposition rates by 30% and increased cellulose decomposition by 9%. In the same set of litter decomposition studies increased exogenous N availability increased the amount of litter entering slowly decaying humus pools in a manner significantly related to the lignocellulose index of decaying litter. Incorporating changes to decomposition rates in TRACE did not accurately reproduce greater soil C storage observed in our field study with experimentally elevated N deposition. However, when changes in the extent of decomposition were incorporated in TRACE, the model produced increased soil C storage by increasing the amount of litter entering the humus pool and accurately represented C storage in plant and soil pools under experimental N deposition. Our modeling results and meta-analysis indicate that the extent of litter decay as humus is formed, rather than slower rates of litter decay, is likely responsible for the accumulation of organic matter, and hence soil C storage, under experimental N deposition. This effect should be incorporated in regional to global-scale models simulating the C balance of forest ecosystems in regions receiving elevated N deposition.     

Does nitrogen availability control rates of litter decomposition in forests?

The effects of increased exogenous N availability on rates of litter decomposition were assessed in several field fertilization trials. In a jack pine (Pinus banksiana Lamb.) forest, needle litter decomposed at the same rate in control plots and in plots fertilized with urea and ammonium nitrate (1350 kg N ha^-1) with or without P and K. Mixed needle litter of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western red cedar (Thuja plicata Donn) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) incubated in plots recently amended with sewage sludge (500 kg N ha^-1) lost less weight during 3 years than did litter in control plots. Forest floor material also decomposed more slowly in plots amended with sewage sludge. Paper birch (Betula papyrifera Marsh.) leaf litter placed on sewage sludge (1000 kg N ha^-1), pulp sludge, or sewage-pulp sludge mixtures decomposed at the same rate as leaf litter in control plots. These experiments demonstrate little effect of exogenous N availability on rates of litter decomposition.The influence of endogenous N availability on rates of litter decomposition was examined in a microcosm experiment. Lodgepole pine (Pinus contorta var. latifolia Engelm.) needle litter collected from N-fertilized trees (525 kg N ha^-1 in ammonium nitrate) were 5 times richer in N than needles from control trees (1.56% N versus 0.33% N in control trees), but decomposed at the same rate. Green needles from fertilized trees contained twice as much N as needles from control trees (1.91% N versus 0.88% N), but decomposed at the same rate. These experiments suggest that N availability alone, either exogenous or endogenous, does not control rates of litter decomposition. Increased N availability, through fertilization or deposition, in the absence of changes in vegetation composition, will not alter rates of litter decomposition in forests.     

Effect of nitrogen addition on decomposition of Calamagrostis angustifolia litters from freshwater marshes of Northeast China

Wetland ecosystems store a large amount of organic carbon (C) in soils, due to the slow decomposition rates of plant litter and soil organic matter. Increased nitrogen (N) availability induced by human activities and global warming may accelerate litter decomposition and affect soil organic C dynamics in wetlands. In the present study, we investigated the effect of N addition on decomposition of Calamagrostis angustifolia litters from freshwater marshes in the Sanjiang Plain of Northeast China under field and laboratory conditions. First, we assessed the changes in initial litter chemical composition and subsequent decomposition following three years of N addition at the rate of 24 g N m⁻² year⁻¹ under field conditions. Our results showed that N addition increased litter N concentration and decreased C/N ratio, and thus stimulated litter decomposition. Secondly, we examined the effect of increased N availability (0, 25, 50 and 100 mg N g⁻¹ litter) on litter decomposition under laboratory conditions. Increased exterior N availability also enhanced microbial respiration and increased litter mass loss under both waterlogging and non-waterlogging conditions. In addition, waterlogging conditions inhibited microbial respiration and suppressed litter mass loss. These findings demonstrated that N addition increased litter decomposition rates through improved litter quality and enhanced microbial activity in freshwater marshes of Northeast China. This implies that increased N availability accelerates litter decomposition rates, and thus may cause substantial losses of soil C and diminish and even reverse the C sink function of wetlands in the Sanjiang Plain of Northeast China.