Effects of Long-Term Nitrogen Addition on Microbial Enzyme Activity in Eight Forested and Grassland Sites: Implications for Litter and Soil Organic Matter Decomposition

Long-term nitrogen (N) addition experiments have found positive, negative, and neutral effects of added N on rates of decomposition. A leading explanation for this variation is differential effects of N on the activity of microbially produced extracellular enzymes involved in decomposition. Specifically, it is hypothesized that adding N to N-limited ecosystems increases activity of cellulose degrading enzymes and decreases that of lignin degrading enzymes, and that shifts in enzyme activity in response to added N explain the decomposition response to N fertilization. We measured litter and soil organic matter (SOM) decomposition and microbial enzyme activity in a long-term N fertilization experiment at eight forested and grassland sites in central Minnesota, USA, to determine (1) variation among sites in enzyme activity, (2) variation in the response of enzymes, litter decomposition, and soil respiration to added N, and (3) whether changes in enzyme activity in response to added N explained variability among sites in the effect of N on litter and SOM decomposition. Site differences in pH, moisture, soil carbon, and microbial biomass explained much of the among-site variation in enzyme activity. Added N generally stimulated activities of cellulose degrading and N- and phosphorus-acquiring enzymes in litter and soil, but had no effect on lignin degrading enzyme activity. In contrast, added N generally had negative or neutral effects on litter and SOM decomposition in the same sites, with no correspondence between effects of N on enzyme activity and decomposition across sites. B.L.K. and S.E.H. conceived of study; B.L.K., S.E.H., and L.E.K. designed study and performed research; B.L.K. analyzed data and wrote the paper.     

N2-Fixing Red Alder Indirectly Accelerates Ecosystem Nitrogen Cycling

Symbiotic N₂-fixing tree species can accelerate ecosystem N dynamics through decomposition feedbacks via both direct and indirect pathways. Direct pathways include the production of readily decomposed leaf litter and increased N supply to decomposers, whereas indirect pathways include increased tissue N and altered detrital dynamics of non-fixing vegetation. To evaluate the relative importance of direct and indirect pathways, we compared 3-year decomposition and N dynamics of N₂-fixing red alder leaf litter (2.34% N) to both low-N (0.68% N) and high-N (1.21% N) litter of non-fixing Douglas-fir, and decomposed each litter source in four forests dominated by either red alder or Douglas-fir. We also used experimental N fertilization of decomposition plots to assess elevated N availability as a potential mechanism of N₂-fixer effects on litter mass loss and N dynamics. Direct effects of N₂-fixing red alder on decomposition occurred primarily as faster N release from red alder than Douglas-fir litter. Direct increases in N supply to decomposers via experimental N fertilization did not stimulate decomposition of either species litter. Fixed N indirectly influenced detrital dynamics by increasing Douglas-fir tissue and litter N concentrations, which accelerated litter N release without accelerating mass loss. By increasing soil N, tissue N, and the rate of N release from litter of non-fixers, we conclude that N₂-fixing vegetation can indirectly foster plant–soil feedbacks that contribute to the persistence of elevated N availability in terrestrial ecosystems.     

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.     

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.     

Regional Contingencies in the Relationship between Aboveground Biomass and Litter in the World’s Grasslands

Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex.     

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.     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

Global decomposition experiment shows soil animal impacts on decomposition are climate‐dependent

Climate and litter quality are primary drivers of terrestrial decomposition and, based on evidence from multisite experiments at regional and global scales, are universally factored into global decomposition models. In contrast, soil animals are considered key regulators of decomposition at local scales but their role at larger scales is unresolved. Soil animals are consequently excluded from global models of organic mineralization processes. Incomplete assessment of the roles of soil animals stems from the difficulties of manipulating invertebrate animals experimentally across large geographic gradients. This is compounded by deficient or inconsistent taxonomy. We report a global decomposition experiment to assess the importance of soil animals in C mineralization, in which a common grass litter substrate was exposed to natural decomposition in either control or reduced animal treatments across 30 sites distributed from 43°S to 68°N on six continents. Animals in the mesofaunal size range were recovered from the litter by Tullgren extraction and identified to common specifications, mostly at the ordinal level. The design of the trials enabled faunal contribution to be evaluated against abiotic parameters between sites. Soil animals increase decomposition rates in temperate and wet tropical climates, but have neutral effects where temperature or moisture constrain biological activity. Our findings highlight that faunal influences on decomposition are dependent on prevailing climatic conditions. We conclude that (1) inclusion of soil animals will improve the predictive capabilities of region‐ or biome‐scale decomposition models, (2) soil animal influences on decomposition are important at the regional scale when attempting to predict global change scenarios, and (3) the statistical relationship between decomposition rates and climate, at the global scale, is robust against changes in soil faunal abundance and diversity.     

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.     

Browsing-induced Effects on Leaf Litter Quality and Decomposition in a Southern African Savanna

We investigated the linkages between leaf litter quality and decomposability in a savanna plant community dominated by palatable-spinescent tree species. We measured: (1) leaf litter decomposability across five woody species that differ in leaf chemistry; (2) mass decomposition, nitrogen (N); and carbon (C) dynamics in leaf litter of a staple browse species (Acacia nigrescens) as well as (3) variation in litter composition across six sites that experienced very different histories of attack from large herbivores. All decomposition trials included litter bags filled with chopped straw to control for variation in site effects. We found a positive relationship between litter quality and decomposability, but we also found that Acacia and straw litter mass remaining did not significantly vary between heavily and lightly browsed sites. This is despite the fact that both the quality and composition of litter returned to the soil were significantly different across sites. We observed greater N resorption from senescing Acacia leaves at heavily browsed sites, which in turn contributed to increase the C:N ratio of leaf litter and caused greater litter N immobilization over time. This, together with the significantly lower tree- and herb-leaf litter mass beneath heavily browsed trees, should negatively affect decomposition rates. However, estimated dung and urine N deposition from both browsers and grazers was significantly greater at high- than at low-herbivory sites. We hypothesize that N inputs from dung and urine boost litter N mineralization and decomposition (especially following seasonal rainfall events), and thereby offset the effects of poor leaf litter quality at chronically browsed sites. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Positive feedbacks between decomposition and soil nitrogen availability along fertility gradients

Background and aims

We determined the relationship between site N supply and decomposition rates with respect to controls exerted by environment, litter chemistry, and fungal colonization.

Methods

Two reciprocal transplant decomposition experiments were established, one in each of two long-term experiments in oak woodlands in Minnesota, USA: a fire frequency/vegetation gradient, along which soil N availability varies markedly, and a long-term N fertilization experiment. Both experiments used native Quercus ellipsoidalis E.J. Hill and Andropogon gerardii Vitman leaf litter and either root litter or wooden dowels.

Results

Leaf litter decay rates generally increased with soil N availability in both experiments while belowground litter decayed more slowly with increasing soil N. Litter chemistry differed among litter types, and these differences had significant effects on belowground (but not aboveground) decay rates and on aboveground litter N dynamics during decomposition. Fungal colonization of detritus was positively correlated with soil fertility and decay rates.

Conclusions

Higher soil fertility associated with low fire frequency was associated with greater leaf litter production, higher rates of fungal colonization of detritus, more rapid leaf litter decomposition rates, and greater N release in the root litter, all of which likely enhance soil fertility. During decomposition, both greater mass loss and litter N release provide mechanisms through which the plant and decomposer communities provide positive feedbacks to soil fertility as ultimately driven by decreasing fire frequency in N-limited soils and vice versa.     

Asymbiotic Nitrogen Fixation and Litter Decomposition on a Long Soil-Age Gradient in Hawaiian Montane Rain Forest1

We determined rates of decomposition and asymbiotic nitrogen fixation in the leaf litter of Cheirodendron spp. on the Hawaiian Islands. Leaf litter was collected from four sites on a long soil-age gradient (300 yr to 4.1 M yr) and decomposed at two sites that differed widely in substrate age and nutrient availability. Rates of decomposition were higher in litter decomposed at the older site, where nutrient availability was greater. A substantial amount of nitrogen and phosphorus immobilization occurred in litter decomposed at the older site, with more immobilization occurring in litter with lower initial nitrogen and phosphorus concentrations, suggesting both supply and demand controls on nutrient immobilization. Potential rates of nitrogen fixation were very low in the first 25 d (0–5 nmol acetylene/gdw/h), rose to much higher rates by 70 d (20–45 nmol), and then declined by 140 d. We found no significant difference in rates of potential nitrogen fixation between sites of decomposition, but there was a strong substrate effect, with higher rates in litter with low lignin, low nitrogen, and high phosphorus. Where significant immobilization of nitrogen occurred for decomposing Cheirodendron, nitrogen fixation could have comprised no more than 10 percent of immobilized nitrogen. Overall, rates of nitrogen fixation were dependent on the source of the decomposing substrate but not on the site of decomposition, while short-term decomposition and nutrient immobilization were strongly dependent on the site of decomposition but not as much on the source of the decomposing substrate.     

Invasive Grass Alters Litter Decomposition by Influencing Macrodetritivores

The results of nitrogen (N) fertilization experiments have shown inconsistent rates of plant litter decomposition, a phenomenon that may be explained by dispropotionate influence of animal detritivores (macro-detritivores) on litter mass loss versus that of microbial decomposers, whose activity may be dependent on inorganic N. In turn, macrodetritivores may be influenced by plant species composition via their selection of optimal food resources and habitats. In our experiment, fertilizer had no apparent effect on litter decomposition, suggesting that microbial decomposers did not use the additional inorganic N and/or that macrodetritivores had a greater influence on decomposition. Manipulation of macrodetritivores suggested that plant species composition—dominated in this study by Festuca arundinacea, an exotic, invasive grass, and Aster ericoides, a native forb—caused shifts in detrivore communities and/or feeding patterns that tended to increase litter mass loss. Canopy cover of F. arundinacea and A. ericoides ranged from 0% to 11%, suggesting that low-intensity invasion may produce significant changes in ecosystem function, such as decomposition.     

Early stages of root and leaf decomposition in Hawaiian forests: effects of nutrient availability

We examined the effects of soil nutrient availability and tissue chemistry on decomposition of both fine roots (<2 mm diameter) and leaves in three sites along a forest chronosequence in the Hawaiian Islands. These sites form a natural fertility gradient, with the youngest and oldest sites having lower nutrient availability than the intermediate-aged site. Nitrogen (N) limits aboveground net primary productivity (ANPP) in the youngest site, while phosphorus (P) limits ANPP in the oldest site. Both root and leaf litter decomposed most slowly in the 4.1-Myear-old site. We also investigated root decomposition in fertilized plots at the youngest and oldest sites; when roots were produced and decomposed in fertilized plots, root decomposition rates increased with N and P additions at the 4.1-Myear-old site. At the 300-year-old site, however, root decomposition rates did not respond to N or P additions. Roots decomposed faster than leaves at the more infertile sites, in part because of lower lignin-to-nitrogen ratios in roots than in leaf litter. Decomposing roots immobilized more nutrients than did decomposing leaves, and may serve an important role in retaining nutrients in these forests. Received: 30 November 1998 / Accepted: 12 August 1999     

How Phosphorus Availability Affects the Impact of Nitrogen Deposition on <Emphasis Type="Italic">Sphagnum</Emphasis> and Vascular Plants in Bogs

To elucidate the impact of high nitrogen (N) deposition on intact bog vegetation, as affected by phosphorus (P) availability, we conducted a 4-year fertilization experiment with N and P at six sites, one with moderate N deposition and five with high N deposition. During the growing season, N (40 kg ha^–1 ya^–1), P (3 kg ha^–1 y^–1), or a combination of both elements was applied to the vegetation. The fertilization effects turned out to be additive in nature. Adding P decreased the inorganic N concentration and increased the P concentration in the rhizosphere at two sites. Furthermore, P stimulated Sphagnum growth and Sphagnum net primary productivity (NPP) at two sites; it also seemed to encourage growth at two other sites including the moderate-deposition site. Vascular plant growth remained largely unaffected but was depressed at one high-deposition site. Adding N increased the concentration of inorganic N in the rhizosphere at the moderate-deposition site and at two of the three high-deposition sites; vascular plant growth and litter production were encouraged at three high-deposition sites. The addition of N depressed Sphagnum height increment at four high deposition sites and reduced Sphagnum NPP at two sites. Shading by vascular plants was of minor importance in explaining the negative effects of N on Sphagnum. We conclude that because P alleviates the negative impact N has on Sphagnum by enhancing its capability to assimilate the deposited N, P availability is a major factor determining the impact of N deposition on Sphagnum production and thus on carbon sequestration in bogs.     

Distinct Microbial Limitations in Litter and Underlying Soil Revealed by Carbon and Nutrient Fertilization in a Tropical Rainforest

Human-caused alterations of the carbon and nutrient cycles are expected to impact tropical ecosystems in the near future. Here we evaluated how a combined change in carbon (C), nitrogen (N) and phosphorus (P) availability affects soil and litter microbial respiration and litter decomposition in an undisturbed Amazonian rainforest in French Guiana. In a fully factorial C (as cellulose), N (as urea), and P (as phosphate) fertilization experiment we analyzed a total of 540 litterbag-soil pairs after a 158-day exposure in the field. Rates of substrate-induced respiration (SIR) measured in litter and litter mass loss were similarly affected by fertilization showing the strongest stimulation when N and P were added simultaneously. The stimulating NP effect on litter SIR increased considerably with increasing initial dissolved organic carbon (DOC) concentrations in litter, suggesting that the combined availability of N, P, and a labile C source has a particularly strong effect on microbial activity. Cellulose fertilization, however, did not further stimulate the NP effect. In contrast to litter SIR and litter mass loss, soil SIR was reduced with N fertilization and showed only a positive effect in response to P fertilization that was further enhanced with additional C fertilization. Our data suggest that increased nutrient enrichment in the studied Amazonian rainforest can considerably change microbial activity and litter decomposition, and that these effects differ between the litter layer and the underlying soil. Any resulting change in relative C and nutrient fluxes between the litter layer and the soil can have important consequences for biogeochemical cycles in tropical forest ecosystems.     

氮、磷养分有效性对森林凋落物分解的影响研究进展

通过对相关研究文献的综述结果表明,氮(N)和磷(P)是构成蛋白质和遗传物质的两种重要组成元素,限制森林生产力和其他生态系统过程,对凋落物分解产生深刻影响。大量的凋落物分解试验发现在土壤N有效性较低的温带和北方森林,凋落物分解速率常与底物初始N浓度、木质素/N比等有很好的相关关系,也受外源N输入的影响;而在土壤高度风化的热带亚热带森林生态系统中,P可能是比N更为重要的分解限制因子。然而控制试验表明,N、P添加对凋落物分解速率的影响并不一致,既有促进效应也有抑制效应。为了深入揭示N、P养分有效性对凋落物分解的调控机制,"底物的C、N化学计量学"假说、"微生物的N开采"假说以及养分平衡的理论都常被用于解释凋落物分解速率的变化。由于微生物分解者具有较为稳定的C、N、P等养分需求比例,在不同的养分供应的周围环境中会体现出不同的活性,某种最缺乏的养分可能就是分解的最重要限制因子。未来的凋落物分解研究,应延长实验时间、加强室内和野外不同条件下的N、P等养分添加控制试验,探讨驱动分解进程的微生物群落结构和酶活性的变化。     

Nitrogen limitation of decomposition and decay: How can it occur?

The availability of nitrogen (N) is a critical control on the cycling and storage of soil carbon (C). Yet, there are conflicting conceptual models to explain how N availability influences the decomposition of organic matter by soil microbial communities. Several lines of evidence suggest that N availability limits decomposition; the earliest stages of leaf litter decay are associated with a net import of N from the soil environment, and both observations and models show that high N organic matter decomposes more rapidly. In direct contrast to these findings, experimental additions of inorganic N to soils broadly show a suppression of microbial activity, which is inconsistent with N limitation of decomposition. Resolving this apparent contradiction is critical to representing nutrient dynamics in predictive ecosystem models under a multitude of global change factors that alter soil N availability. Here, we propose a new conceptual framework, the Carbon, Acidity, and Mineral Protection hypothesis, to understand the effects of N availability on soil C cycling and storage and explore the predictions of this framework with a mathematical model. Our model simulations demonstrate that N addition can have opposing effects on separate soil C pools (particulate and mineral‐protected carbon) because they are differentially affected by microbial biomass growth. Moreover, changes in N availability are frequently linked to shifts in soil pH or osmotic stress, which can independently affect microbial biomass dynamics and mask N stimulation of microbial activity. Thus, the net effect of N addition on soil C is dependent upon interactions among microbial physiology, soil mineralogy, and soil acidity. We believe that our synthesis provides a broadly applicable conceptual framework to understand and predict the effect of changes in soil N availability on ecosystem C cycling under global change.     

Nitrogen supply differentially affects litter decomposition rates and nitrogen dynamics of sub-arctic bog species

High-latitude peatlands are important soil carbon sinks. In these ecosystems, the mineralization of carbon and nitrogen are constrained by low temperatures and low nutrient concentrations in plant litter and soil organic matter. Global warming is predicted to increase soil N availability for plants at high-latitude sites. We applied N fertilizer as an experimental analogue for this increase. In a three-year field experiment we studied N fertilization effects on leaf litter decomposition and N dynamics of the four dominant plant species (comprising >75% of total aboveground biomass) in a sub-arctic bog in northern Sweden. The species were Empetrum nigrum (evergreen shrub), Eriophorum vaginatum (graminoid), Betula nana (deciduous shrub) and Rubus chamaemorus (perennial forb). In the controls, litter mass loss rates increased in the order: Empetrum < Eriophorum < Betula < Rubus. Increased N availability had variable, species-specific effects: litter mass loss rates (expressed per unit litter mass) increased in Empetrum, did not change in Eriophorum and Betula and decreased in Rubus. In the leaf litter from the controls, we measured no or only slight net N mineralization even after three years. In the N-fertilized treatments we found strong net N immobilization, especially in Eriophorum and Betula. This suggests that, probably owing to substantial chemical and/or microbial immobilization, additional N supply does not increase the rate of N cycling for at least the first three years.     

Effects of mixing pine and broadleaved tree/shrub litter on decomposition and N dynamics in laboratory microcosms

This study was carried out to compare the ecological function of exotic pine (Pinus radiata—Pr) and native pine (Pinus tabulaeformis—Pt) in terms of litter decomposition and its related N dynamics and to evaluate if the presence of broad-leaved tree species (Cercidiphyllum japonicum—Cj) or shrub species (Ostryopsis davidiana—Od) litter would promote the decomposition of pine needles and N cycling. Mass remaining, N release of the four single-species litters and mixed-species (Pt + Cj; Pr + Cj; Pt + Od; Pr + Od) litters and soil N dynamics were measured at microcosm scale during an 84-day incubation period. The Pt and Pr litter, with poorer substrate quality, indicated slower decomposition rates than did the Cj and Od litter. Due to their high C/N ratios, the N mass of Pt and Pr litter continuously increased during the early stage of decomposition, which showed that Pt and Pr litter immobilized exogenous N by microbes. No significant differences of soil inorganic, dissolved organic and microbial biomass N were found between the Pt and Pr microcosm at each sampling. The results showed that the exotic Pr performed similar ecological function to the native Pt in terms of litter decomposition and N dynamics during the early stage. The presence of Cj or Od litter increased the decomposition rates of pine needle litter and also dramatically increased soil N availability. So it is feasible for plantation managers to consider the use of Cj as an ameliorative species or to retain Od in pine plantations to promote the decomposition of pine litter and increase nutrient circulation. The results also suggested that different species litters induced different soil dissolved organic nitrogen (DON). As a major soluble N pool in soil, DON developed a different changing tendency over time compared with inorganic N, and should be included into soil N dynamic under the condition of our study.