Nitrogenase activity on the roots of tropical forage grasses

Nitrogen fixation in the rhizospheres of field grown tropical forage grasses was studied by the acetylene reduction method. Values varied considerably between sites but indicate the possible economic importance of several of the species studied. Maximal nitrogenase activity measured (nmoles C₂H₄g^?1 dry roots h^?1) was 754 for Pennisetum purpureum, 750 for Brachiaria mutica, 341 for Digitaria decumbens, 299 for Panicum maximum, 283 for Paspalum notatum, 269 for Cynodon dactylon, 41 for Melinis minutiflora and 29 for Hyparrhenia rufa. Nitrogenase activity varied considerably with season and was maximal during active vegetative growth of two of the grasses. Significant differences between Paspalum notatum ecotypes and cultivars. in Azotohacter paspali occurrence and nitrogen fixation, indicate the possibility of plant breeding to enhance nitrogen fixation in grass rhizospherc associations. Other research lines of agronomic importance are fertilizer effects. In intact soil plant cores with the Paspalum system 10 parts/10⁶ NH_4⁺J-N inhibited nitrogenase activity within 2 h and 10 parts/10⁶ NO^?₃-N within 4 h. but after 1 week these effects were negligible. In the field, nitrogenase activity on roots of P. purpureum and D. decumbens, assayed 2 weeks after top dressings of 20 kg N ha^?1 as NH₄NO₃. was not affected even after eight such dressings.     

Nitrogen availability mediates the effects of roots and mycorrhizal fungi on soil organic carbon decomposition: A meta-analysis

Plant roots and their associated mycorrhizal fungi critically mediate the decomposition of soil organic carbon (C), but the general patterns of their impacts over a broad geographical range and the primary mediating factors remain unclear. Based on a synthesis of 596 paired observations from both field and greenhouse experiments, we found that living roots and/or mycorrhizal fungi increased organic C decomposition by 30.9%, but low soil nitrogen (N) availability (i.e., high soil C:N ratio) critically mitigated this promotion effect. In addition, the positive effects of living roots and/or mycorrhizal fungi on organic C decomposition were higher under herbaceous and leguminous plants than under woody and non-leguminous plants, respectively. Surprisingly, there was no significant difference between arbuscular mycorrhizal fungi and ectomycorrhizal fungi in their effects on organic C decomposition. Furthermore, roots and/or mycorrhizal fungi significantly enhanced the decomposition of leaf litter but not root litter. These findings advance our understanding of how roots and their symbiotic fungi modulate soil C dynamics in the rhizosphere or mycorrhizosphere and may help improve predictions of soil global C balance under a changing climate.     

Effect of earthworm addition on soil nitrogen availability,microbial biomass and litter decomposition in mesocosms

The aim of the study was to determine the effect of adding two tropical earthworm species, Rhinodrilus contortus and Pontoscolex corethrurus, to mesocosms on the availability of mineral N (NH₄ ⁺ and NO₃ ^– concentrations), soil microbial biomass (bio-N), and the decomposition rates of three contrasting leaf litter species, in a glasshouse experiment. The mesocosms were filled with forest soil and covered with a layer of leaf litter differing in nutritional quality: (1) Hevea brasiliensis (C/N=27); (2) Carapa guianensis (C/N=32); (3) Vismia sp., the dominant tree species in the second growth forest (control, C/N= 42); and, (4) a mixture of the former three leaf species, in equal proportions (C/N=34). At the end of the 97-day experiment, the soil mineral N concentrations, bio-N, and leaf litter weight loss were determined. Both earthworm species showed significant effects on the concentrations of soil NO₃ ^– (p<0.01) and NH₄ ⁺ (p<0.05). Bio-N was always greater in the mesocosms with earthworms (especially with R. contortus) and in the mesocosms with leaf litter of H. brasiliensis (6 µg N g^–1 soil), the faster decomposing species, than in the other treatments (0.1–1.6 µg N g^–1). Thus, earthworm activity increased soil mineral-N concentrations, possibly due to the consumption of soil microbial biomass, which can speed turnover and mineralization of microbial tissues. No significant differences in decomposition rate were found between the mesocosms with and without earthworms, suggesting that experiments lasting longer are needed to determine the effect of earthworms on litter decomposition rates.     

Direct effects of litter decomposition on soil dissolved organic carbon and nitrogen in a tropical rainforest

To clarify how litter decomposition processes affect soil dissolved organic carbon (DOC) and soil dissolved nitrogen (DN) dynamics, we conducted a field experiment on leaf litter and collected DOC and DN from the underlying soil in a tropical rainforest in Xishuangbanna, southwest China. Principal components analysis (PCA) showed the first PCA axis (corresponding to degraded litter quantity and quality) explained 61.3% and 71.2% of variation in DOC and DN concentrations, respectively. Stepwise linear regression analysis indicated that litter carbon mass controlled DOC and hemicellulose mass controlled DN concentrations. Litter decomposition was the predominant factor controlling surface-soil DOC and DN dynamics in this tropical rainforest.     

Effect of active roots on the decomposition of soil organic materials

Summary The effect of one form of soil organic matter, such as living roots or root exudates on another form of soil organic matter, such as dead roots or incorporated litter and litter leachates, has been studied from various perspectives over the last 25 years. The effect seems to be either positive (priming) or negative (conserving). The present review concentrates on the conserving effect, measured as a decrease in ¹⁴CO₂ released, in both field and greenhouse/growth chamber studies. The field experiments suggested that certain physical conditions in the soil, such as less available moisture or restricted aeration which led to lower microbial activity, explained the conserving effect of living roots on soil organic matter. Although more detailed greenhouse/growth chamber studies confirmed the conserving effect per se, it appears that biological rather than physical factors could better explain the reduction in the rate of decomposition of ¹⁴C-labelled plant residues in the presence of roots. However, a complex picture has emerged through a variety of postulates, all proposed in attempts to explain the conserving effect. Finally, the most recent studies have argued that the decrease in decomposition of labelled organic matter in planted soil is probably more apparent than real. A decrease in respired ¹⁴CO₂ could be explained by an incorporation of ¹⁴C derived from old roots into the rhizosphere microbial populations of the living roots. To make any further progress on the fundamental question of how soil organic matter moves along its continuum from a living to a refractory state, the microenvironment needs to be examined at periodic intervals. New developments in improved histochemical and electron-probe microanalyses look promising.LRS Contribution no. 3878970     

长期氮肥施用对玉米根茬矿化分解的影响

通过室内培养和田间分解试验,研究了施氮量为N 0、120和240 kg/hm2处理的玉米根茬(R0、R120、R240)在15和45 cm两个肥力条件不同的土层中有机碳矿化分解特性及其对土壤活性有机碳组分的影响。结果表明,在室内矿化培养条件下,根茬CO2累积释放量和潜在碳矿化量均为R120R240R0;R120和R240根茬碳矿化率在表层土壤(15 cm)和底层土壤(45 cm)中分别较R0提高21.1%、12.7%和45.3%、33.7%。在田间埋藏分解条件下,分解386 d后R0、R120和R240根茬碳残留率在表层土壤中分别为36.3%、25.2%和28.7%,在底层土壤中分别为38.4%、30.6%和31.1%;根茬碳残留率与其C/N、木质素含量以及木质素/N正相关,而与根茬全氮含量呈负相关关系,表明根茬分解率随着其本身全氮含量的增加而提高;添加玉米根茬显著增加土壤微生物量碳含量143%~297%,增加土壤可溶性有机碳含量19.9%~118.2%。综上可见,长期施用氮肥影响作物根系的养分组成,显著提高其全氮含量,在评价土壤碳、氮养分循环时,应注重长期氮肥施用对作物残茬养分累积及其在土壤中分解、转化的影响。     

Soil characteristics determine soil carbon and nitrogen availability during leaf litter decomposition regardless of litter quality

Climate and litter quality have been identified as major drivers of litter decomposition, but our knowledge of how soil characteristics (e.g. microbial community and chemical properties) determine carbon (C) and nitrogen (N) availability derived from the decomposition of litter of different qualities is still scarce. We conducted a microcosm experiment to evaluate how soils with contrasting microbial communities and soil properties (denoted Soils A and B hereafter, where Soil B has higher bacterial and fungal abundance, fungal:bacterial ratio, and organic C than Soil A) determine the availability of soil C (carbohydrates, proteins, amino acids and phenols) and N (dissolved organic and inorganic N, microbial biomass N and available N) during the decomposition of litter of contrasting quality (C:N ratios ranging from 20 to 102). We also evaluated the relative importance of soil characteristics and litter quality as drivers of C and N inputs to the soil during this process. Overall, higher soil C and N availability after litter decomposition was found in Soil B than in Soil A. Soil characteristics had a higher positive effect on soil C and N contents than litter quality during litter decomposition. We also found that changes in N availability and organic matter quality registered after litter decomposition, linked to different soil characteristics, were able to promote dissimilarities in the potential mineralization rates. In conclusion, our study provides evidence that soil characteristics (e.g. microbial communities and chemical properties) can be more important than litter quality in determining soil C and equally important for N availability during the decomposition of leaf litter.     

Influence of cover crops on potential nitrogen availability to succeeding crops in a Southern Piedmont soil

Winter cover crops are essential in conservation tillage systems to protect soils from erosion and for improving soil productivity. Black oat (Avena strigosa Schreb) and oilseed radish (Raphanus sativus L.) could be useful cover crops in the southeastern USA, but successful adoption requires understanding their influence on N availability in conservation tillage systems. Black oat and oilseed radish were compared to crimson clover (Trifolium incarnatum L.) and rye (Secale cereale L.) for biomass production and effects on N mineralization during the summer crop growing season from fall 1998 through summer 2002 near Watkinsville, GA. Rye produced 40 to 60% more biomass, although N contents were less than the other cover crops. Oilseed radish and black oat N contents were similar to crimson clover. Black oat, oilseed radish, and crimson clover C/N ratios were less than 30, whereas rye averaged 39. Amount of N mineralized in 90 days (N _min90) measured with in situ soil cores was 1.3 to 2.2 times greater following black oat, crimson clover, and oilseed radish than following rye. No differences in N _min90 were found between black oats, crimson clover, and oilseed radish in 1999 and 2000. The amount of potentially mineralizable N (N ₀) was not different due to cover crop, but was 1.5 times greater in 2000 and 2002 than in 1999. The rate of N mineralization (k) was 20 to 50% slower following rye than the other three cover crops. Black oat and oilseed radish biomass production and soil N mineralization dynamics were more similar to crimson clover than to rye, which indicates that they could be used as cover crops in the southeast without significant changes in N recommendations for most crops.

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Effects of pine roots on microorganisms,fauna, and nitrogen availability in two soil horizons of a coniferous forest spodosol

Summary We investigated the effects of pitch pine seedling roots on extractable N, microbial growth rate, biomass C and N, and nematodes and microarthropods in microcosms with either organic (41% C, 1.14% N) or mineral (0.05% C, 0.01% N) horizon soils of a spondosol. Root quantity was manipulated by varying plant density (0, 1, 2, or 4 seedlings) and rhizosphere soil was separated from non-rhizosphere soil by a 1.2 m mesh fabric. In the rhizosphere of organic soil horizons, moisture, microbial growth rate, biomass C and N, and extractable N declined as root density was increased, but there was little effect on nematodes or microarthropods. High levels of extractable N remained after 5 months, suggesting that N mineralization was stimulated during the incubation. In the rhizosphere of mineral soil horizons, microbial growth rate, and nematode and microarthropod abundances increased at higher root density, and in the absence of roots faunal abundance approached zero. Faunal activity was concentrated in the rhizosphere compared to non-rhizosphere soil. In organic soil horizons, roots may limit microbial activity by reducing soil moisture and/or N availability. However, in mineral soil horizons, where nutrient levels are very low, root inputs can stimulate microbial growth and faunal abundance by providing important substrates for microbial growth. Our results demonstrate a rhizosphere effect for soil fauna in the mineral soil, and thus extends the rhizosphere concept to components of the soil community other than microbes for forest ecosystems. Although our results need to be verified by field manipulations, we suggest that the effects of pine roots on nutrient cycling processes in coniferous forests can vary with soil nutrient content and, therefore, position in the soil profile.     

N2O emissions from humid tropical agricultural soils: effects of soil moisture,texture and nitrogen availability

We studied soil moisture dynamics and nitrous oxide (N₂O) fluxes from agricultural soils in the humid tropics of Costa Rica. Using a split-plot design on two soils (clay, loam) we compared two crop types (annual, perennial) each unfertilized and fertilized. Both soils are of andic origin. Their properties include relatively low bulk density and high organic matter content, water retention capacity, and hydraulic conductivity. The top 2–3 cm of the soils consists of distinct small aggregates (dia. <0.5 cm). We measured a strong gradient of bulk density and moisture within the top 7 cm of the clay soil. Using automated sampling and analysis systems we measured N₂O emissions at 4.6 h intervals, meteorological variables, soil moisture, and temperature at 0.5 h intervals. Mean daily soil moisture content at 5 cm depth ranged from 46% water filled pore space (WFPS) on clay in April 1995 to near saturation on loam during a wet period in February 1996. On both soils the aggregated surface layer always remained unsaturated. Soils emitted N₂O throughout the year. Mean N₂O fluxes were 1.04±0.72 ng N₂O-N cm⁻² h⁻¹ (mean±standard deviation) from unfertilized loam under annual crops compared to 3.54±4.31 ng N₂O-N cm⁻² h⁻¹ from the fertilized plot (351 days measurement). Fertilization dominated the temporal variation of N₂O emissions. Generally fluxes peaked shortly after fertilization and were increased for up to 6 weeks (‘post fertilization flux’). Emissions continued at a lower rate (‘background flux’) after fertilization effects faded. Mean post-fertilization fluxes were 6.3±6.5 ng N₂O-N cm⁻² h⁻¹ while the background flux rate was 2.2±1.8 ng N₂O-N cm⁻² h⁻¹. Soil moisture dynamics affected N₂O emissions. Post fertilization fluxes were highest from wet soils; fluxes from relatively dry soils increased only after rain events. N₂O emissions were weakly affected by soil moisture during phases of low N availability. Statistical modeling confirmed N availability and soil moisture as the major controls on N₂O flux. Our data suggest that small-scale differences in soil structure and moisture content cause very different biogeochemical environments within the top 7 cm of soils, which is important for net N₂O fluxes from soils.     

Influence of cultivation on soil nitrogen pools

Abstract

Exchangeable NH₄, organic N, and fixed NH₄, were followed in three soil layers (0–25, 25–50, and 50–75 cm) of plots under conventional and minimum tillage in a 10—year field experiment. The main effect of both tillage treatments was a marked increase of fixed NH₄ during the first two years which was attributed to the heavy application of N fertilizers because soils were not fertilized prior to the experiment. Due to spatial variability of soil composition, a statistically significant increase over the 10—year was observed for total and fixed NH₄ only in the surface layer of conventionally tilled soils, probably due to thorough mixing caused by intense cultivation. In this layer the organic N pool did not appear to vary with the years, while the fixed NH₄ pool was influenced by N fertilization. A general trend was a uniform increase of the ratio between fixed NH₄ and total N. Under conventional tillage, the trend was similar for the three soil layers while the reduced amount of fixed NH₄ present in the upper soil layers (0—25 and 25–50 cm) was assumed to be caused by root absorption. Under minimum tillage, the increase of fixed NH₄ ratio was limited to the first 50 cm of soil, and was less pronounced in the top layer where maximum root accumulation is generally expected to be present. The data support the importance for crops of the fixed NH₄ pool.     

Influence of different land uses on soil nitrogen transformations after conversion from an Indian dry tropical forest

The effects of alternate land uses, such as grassland, cropland and mine spoil on mineral nitrogen (N), N-transformation rate and microbial biomass N (MBN) in dry tropical forest soils of India were studied. The mean annual mineral N in the forest, grassland, cropland and mine spoil ecosystems, respectively ranged from 15.24 to 19.58, 17.8 to 18.56, 16.49 to 19.85 and 10.52 to 13.44 µg g^− 1, net nitrification rate from 14.15 to 23.4, 10.11 to 11.38, 8.07 to 9.16, 10.52 to 13.44 µg g^− 1mo^− 1; net N-mineralization rate from 17.38 to 26.36, 13.99 to 15.41, 10.99 to 12.5, 5.43 to 7.68 µg g^− 1mo^− 1and and microbial biomass N from 41.25 to 58.87, 34.47 to 47.95, 27.88 to 30.43 and 22.95 to 25.26 µg g^− 1, respectively. The values were within the range reported by previous studies in different tropical environments. The mean annual net nitrification rates declined after conversion into grassland, cropland and mine spoil by 43, 54 and 78%, respectively, net N mineralization by 33, 46 and 70%, and microbial biomass N by 29%, 42% and 52%, respectively.The MBN was positively related to root biomass and total plant biomass, while microbial-N and inorganic N are reciprocally, while nitrification and N-mineralization are directly related to seasonal soil moisture and temperature. The microbial biomass N, nitrification and N-mineralization are negatively related to smaller fraction (< 0.1 mm) of the soil. Above- and below-ground biomass also have had their impact on microbial biomass N, and thereby N-mineralization. Thus, in dry tropical forests, land-use change affects remarkably the nitrogen transformation process in soil.     

Relating the biological stability of soil organic matter to energy availability in deep tropical soil profiles

Tropical subsoils contain large reservoirs of carbon (C), most of which is stored in soil organic matter (SOM). Subsoil OM is thought to be particularly stable against microbial decomposition due to various mechanisms and its position in the soil profile, potentially representing a long-term C sink. However, few experiments have explicitly investigated SOM stability and microbial activity across several orders of magnitude of soil C concentrations as a function of soil depth. The objective of this study was to evaluate the biological stability of SOM in the upper 1.4 m of tropical forest soil profiles. We did so by measuring CO₂ evolution during a 90-day laboratory incubation experiment on a sample set that was previously characterized for C and nutrient concentrations and microbial biomass. We concurrently measured the energy content of SOM using differential scanning calorimetry (DSC) as an index of the energy available for microbial metabolism, with the hypothesis that the biological stability of SOM would be inversely related to the energy contained within it. Cumulative CO₂ evolution, mean respiration rates, and the energy density of SOM (energy released during combustion normalized to soil C) all declined with soil depth (P < 0.01). Biological indices of C stability were well correlated with measures of SOM energy. There was no change in the mean respiration rate as a function of depth when normalized to soil C, and a trend toward increased respiration per-unit microbial biomass (P = 0.07). While reduced microbial respiration in subsoils suggests an increase in the biological stability of SOM, we suggest this is driven principally by concurrent declines in energy availability as measured by DSC and the size of the microbial biomass pool. On a per-unit biomass basis, subsoil OM may be as prone to decomposition and destabilization as surface SOM.     

Effects of nitrogen enrichment on belowground communities in grassland: Relative role of soil nitrogen availability vs. soil acidification

Terrestrial ecosystems worldwide are receiving increasing amounts of biologically reactive nitrogen (N) as a consequence of anthropogenic activities. This intended or unintended fertilization can have a wide range of impacts on the above- and belowground communities. An increase in high N availability has been assumed to be a major mechanism enhancing the abundance of above- and belowground communities. In addition to increasing available N, however, N enrichment causes soil acidification, which may negatively affect above- and belowground communities. The relative importance of increased N availability vs. increased soil acidity for above- and belowground communities in natural ecosystems experiencing N enrichment is unclear. In a 12-year N enrichment experiment in a semi-arid grassland, N enrichment substantially increased both above- and belowground plant biomass mainly via the N availability-induced increase in biomass of perennial rhizome grasses. N enrichment also dramatically suppressed bacterial, fungal, and actinobacteria biomass mainly via the soil acidification pathway (acidification increased concentrations of H⁺ ions and Al³⁺ and decreased concentrations of mineral cations). In addition, N enrichment also suppressed bacterial-, fungal-feeding, and omnivorous + carnivorous nematodes mainly via the soil acidification pathway (acidification reduced nematode food resources and reduced concentrations of mineral cations). The positive effects resulting from the increase in belowground carbon allocation (via increase in quantity and quality of plant production) on belowground communities were outweighed by the negative effects resulting from soil acidification, indicating that N enrichment weakens the linkages between aboveground and belowground components of grassland ecosystems. Our results suggest that N enrichment-induced soil acidification should be included in models that predict biota communities and linkages to carbon and nitrogen cycling in terrestrial ecosystems under future scenarios of N deposition.     

Effect of composted sewage sludge amendment on soil nitrogen and phosphorus availability

Abstract

Municipal sewage sludge previously composted with sawdust (CSS) was applied to an eutric sandy cambisol at rates of 7.5, 15.0, 22.5, and 30 g#lbkg^‐1. Incubation and pot experiments were conducted to evaluate CSS effectiveness on nitrogen (N) and phosphorus (P) soil availability and on plant nutrition. The CSS rates did not increase soil mineral N and had little effect on organic P and on labile forms of P. Efficiency of total applied P was 17% for the soil labile forms and 4.8% for the resin extractable fraction. In contrast, CSS significantly increased hydroxide extractable inorganic P and nonextractable soil P fraction. The major portion of the increment on nonextractable forms was at the expense of HC1 extractable P fraction [calcium (Ca)‐bounded], dominant on the original CSS. Thus, chemical rather than biological reactions lead to the redistribution of CSS‐borne P to more firmly held forms after its application to the soil. Ryegrass dry matter yield, N content, and N uptake did not increase in CSS‐treated soils. Plant P content increased at the second harvest, but the effect was nil in the subsequent harvest. Total P uptake increased from 14.1 to 20.2 mg#lbpot^‐1, but percentage P recovery by ryegrass was modest, averaging 2.5% of the CSS‐borne P. Results suggest that moderate application of CSS to agricultural systems are inadequate for crop growth but may contribute to nutrient recycling without environmental risks related to N and P loss.     

应用15N示踪技术研究土壤水分对氮素有效性的影响

在有防雨设施的试验田里,设置不同的土壤水分处理,应用15N示踪技术研究土壤水分对氮素有效性的影响.试验结果表明:冬小麦对肥料氮素的利用率随土壤水分提高而提高;土壤供应的有效性氮素(A值)在土壤水分由田间持水量的50%提高到60%时出现"跃迁",土壤水分超过田间持水量的60%以后,A值差异不显著,表明土壤氮素有效性对土壤水分存在一个阈值反映.节水、节肥高效的土壤水分下限应控制在土壤田间持水量的60%以上.     

Earthworms,soil mineral nitrogen and forage production in grass-based hayfields

This study was designed to address how earthworm activity influences soil mineral nitrogen (N), plant N uptake and forage yield in grass-based hayfields. Earthworm populations were reduced by applying carbaryl pesticide to the experimental field plots every 2-weeks, effectively eliminating the earthworms for up to 12-weeks from May to August. Grass yields and tissue N concentrations were measured every 2 weeks, and the soil mineral N concentration determined at the final harvest. Reducing earthworm populations for up to 12-weeks did not affect grass yield or N uptake. However, regression analysis showed that plots with undisturbed earthworm populations had higher soil N by 0.8 kg N ha^?1 per week, representing mineralization of about 10 kg N ha^?1 during the 12-week study. This was a fraction of the fertilizer N recommendation (75 kg N ha^?1) for grass-based hayfields in this region. Therefore, the increase in soil mineral N from earthworm activity was small, relative to the N requirements of the hayfield.     

Effect of added nitrogen on plant litter decomposition depends on initial soil carbon and nitrogen stoichiometry

Increasing organic carbon inputs to agricultural soils through the use of pastures or crop residues has been suggested as a means of restoring soil organic carbon lost via anthropogenic activities, such as land use change. However, the decomposition and retention of different plant residues in soil, and how these processes are affected by soil properties and nitrogen fertiliser application, is not fully understood. We evaluated the rate and extent of decomposition of ¹³C-pulse labelled plant material in response to nitrogen addition in four pasture soils of varying physico-chemical characteristics. Microbial respiration of buffel grass (Cenchrus ciliaris L.), wheat (Triticum aestivum L.) and lucerne (Medicago sativa L.) residues was monitored over 365-days. A double exponential model fitted to the data suggested that microbial respiration occurred as an early rapid and a late slow stage. A weighted three-compartment mixing model estimated the decomposition of both soluble and insoluble plant ¹³C (mg C kg⁻¹ soil). Total plant material decomposition followed the alkyl C: O-alkyl C ratio of plant material, as determined by solid-state ¹³C nuclear magnetic resonance spectroscopy. Urea-N addition increased the decomposition of insoluble plant ¹³C in some soils (≤0.1% total nitrogen) but not others (0.3% total nitrogen). Principal components regression analysis indicated that 26% of the variability of plant material decomposition was explained by soil physico-chemical characteristics (P = 0.001), which was primarily described by the C:N ratio. We conclude that plant species with increasing alkyl C: O-alkyl C ratio are better retained as soil organic matter, and that the C:N stoichiometry of soils determines whether N addition leads to increases in soil organic carbon stocks.