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.     

Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

Long-Term Nitrogen Amendment Alters the Diversity and Assemblage of Soil Bacterial Communities in Tallgrass Prairie

Anthropogenic changes are altering the environmental conditions and the biota of ecosystems worldwide. In many temperate grasslands, such as North American tallgrass prairie, these changes include alteration in historically important disturbance regimes (e.g., frequency of fires) and enhanced availability of potentially limiting nutrients, particularly nitrogen. Such anthropogenically-driven changes in the environment are known to elicit substantial changes in plant and consumer communities aboveground, but much less is known about their effects on soil microbial communities. Due to the high diversity of soil microbes and methodological challenges associated with assessing microbial community composition, relatively few studies have addressed specific taxonomic changes underlying microbial community-level responses to different fire regimes or nutrient amendments in tallgrass prairie. We used deep sequencing of the V3 region of the 16S rRNA gene to explore the effects of contrasting fire regimes and nutrient enrichment on soil bacterial communities in a long-term (20 yrs) experiment in native tallgrass prairie in the eastern Central Plains. We focused on responses to nutrient amendments coupled with two extreme fire regimes (annual prescribed spring burning and complete fire exclusion). The dominant bacterial phyla identified were Proteobacteria, Verrucomicrobia, Bacteriodetes, Acidobacteria, Firmicutes, and Actinobacteria and made up 80% of all taxa quantified. Chronic nitrogen enrichment significantly impacted bacterial community diversity and community structure varied according to nitrogen treatment, but not phosphorus enrichment or fire regime. We also found significant responses of individual bacterial groups including Nitrospira and Gammaproteobacteria to long-term nitrogen enrichment. Our results show that soil nitrogen enrichment can significantly alter bacterial community diversity, structure, and individual taxa abundance, which have important implications for both managed and natural grassland ecosystems.     

Conservation of nitrogen increases with precipitation across a major grassland gradient in the Central Great Plains of North America

Regional analyses and biogeochemical models predict that ecosystem N pools and N cycling rates must increase from the semi-arid shortgrass steppe to the sub-humid tallgrass prairie of the Central Great Plains, yet few field data exist to evaluate these predictions. In this paper, we measured rates of net N mineralization, N in above- and belowground primary production, total soil organic matter N pools, soil inorganic N pools and capture in resin bags, decomposition rates, foliar ¹⁵N, and N use efficiency (NUE) across a precipitation gradient. We found that net N mineralization did not increase across the gradient, despite more N generally being found in plant production, suggesting higher N uptake, in the wetter areas. NUE of plants increased with precipitation, and δ¹⁵N foliar values and resin-captured N in soils decreased, all of which are consistent with the hypothesis that N cycling is tighter at the wet end of the gradient. Litter decomposition appeared to play a role in maintaining this regional N cycling trend: litter decomposed more slowly and released less N at the wet end of the gradient. These results suggest that immobilization of N within the plant–soil system increases from semi-arid shortgrass steppe to sub-humid tallgrass prairie. Despite the fact that N pools increase along a bio-climatic gradient from shortgrass steppe to mixed grass and tallgrass prairie, this element becomes relatively more limiting and is therefore more tightly conserved at the wettest end of the gradient. Similar to findings from forested systems, our results suggest that grassland N cycling becomes more open to N loss with increasing aridity.     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

Progressive N limitation of plant response to elevated CO2: a microbiological perspective

A major uncertainty in predicting long-term ecosystem C balance is whether stimulation of net primary production will be sustained in future atmospheric CO₂ scenarios. Immobilization of nutrients (N in particular) in plant biomass and soil organic matter (SOM) provides negative feedbacks to plant growth and may lead to progressive N limitation (PNL) of plant response to CO₂ enrichment. Soil microbes mediate N availability to plants by controlling litter decomposition and N transformations as well as dominating biological N fixation. CO₂-induced changes in C inputs, plant nutrient demand and water use efficiency often have interactive and contrasting effects on microbes and microbially mediated N processes. One critical question is whether CO₂-induced N accumulation in plant biomass and SOM will result in N limitation of microbes and subsequently cause them to obtain N from alternative sources or to alter the ecosystem N balance. We reviewed the experimental results that examined elevated CO₂ effects on microbial parameters, focusing on those published since 2000. These results in general show that increased C inputs dominate the CO₂ impact on microbes, microbial activities and their subsequent controls over ecosystem N dynamics, potentially enhancing microbial N acquisition and ecosystem N retention. We reason that microbial mediation of N availability for plants under future CO₂ scenarios will strongly depend on the initial ecosystem N status, and the nature and magnitude of external N inputs. Consequently, microbial processes that exert critical controls over long-term N availability for plants would be ecosystem-specific. The challenge remains to quantify CO₂-induced changes in these processes, and to extrapolate the results from short-term studies with step-up CO₂ increases to native ecosystems that are already experiencing gradual changes in the CO₂ concentration.     

Nitrogen uptake and nitrogen use efficiency above and below ground along a topographic gradient of soil nitrogen availability

Nitrogen (N) uptake and nitrogen use efficiency (NUE) are closely related through feedback mechanisms to soil N availability and N cycling in forested ecosystems. We investigated N uptake and NUE not only at the leaf, litterfall, and aboveground levels but also belowground and whole stand levels along a topographic gradient of soil N availability in a cool temperate deciduous forest in Japan. In this study, we addressed how whole stand level N uptake and NUE affect C and N cycling in forested ecosystems. At the leaf, litterfall, and aboveground levels, N uptake decreased and NUE increased with decreasing soil N availability. This pattern resulted from decreasing leaf N concentrations and increasing N resorption efficiencies as soil N availability declined. Low N concentrations in litterfall may have resulted in little soil N being available to plants, due to microbial immobilization. In contrast, when belowground components were included, N uptake and NUE were not correlated with soil N availability. This was mainly due to higher levels of fine root production when soil N availability was low. Higher fine root allocation can result in a high input of detritus to decomposer systems and, thus, contribute to accumulation of soil organic matter and immobilization by microbes, which may result in further soil N availability decline. Our results suggest that allocation to the fine root rather than whole stand level NUE is important for C and N cycling in forested ecosystems, as is the feedback mechanism in which litterfall level NUE shifts with changes in the N concentration of litterfall.     

Soil physiochemical changes following 12 years of annual burning in a humid–subtropical tallgrass prairie: a hypothesis

Burning is known to stimulate growth of grassland vegetation, promote species diversity, and inhibit natural invasion by woody plants. However, the frequency at which grasslands are burned as part of their management can affect soil nutrient content and, ultimately, productivity. The objective of this study was to characterize changes in soil physical and chemical properties in a native tallgrass prairie after 12 years of annual burning. In 1989, five soil samples from the 0 to 10 cm depth were collected along a transect through a 3 ha parcel of native tallgrass prairie in central Arkansas. Soil sampling was repeated in 2001 to assess changes over time. Results showed that soil bulk density, electrical conductivity, extractable P, Na, Fe, and Mn decreased significantly (P < 0.05), while soil organic matter, total N and C, and the C/N ratio increased significantly (P < 0.05) within the 12-year period during which annual burning was the only imposed management practice. Mean extractable K, Ca, Mg, S, and Zn levels were all lower in 2001 than in 1989, but differences were not significant, while soil pH did not change. The results of this study indicate that annual export of several essential plant nutrients during prescribed burning of relatively small, remnant prairie fragments exceeds annual imports from atmospheric deposition and/or organic matter mineralization. Annual prescribed burning may be too frequent to maintain optimal ecosystem functioning and productivity. Decreasing the frequency of prescribed burning for native grassland management may help to retain more soil nutrients to sustain a higher level of productivity.     

Biomass production in a nitrogen-fertilized,tallgrass prairie ecosystem exposed to ambient and elevated levels of CO2

Increased biomass production in terrestrial ecosystems with elevated atmospheric CO₂ may be constrained by nutrient limitations as a result of increased requirement or reduced availability caused by reduced turnover rates of nutrients. To determine the short-term impact of nitrogen (N) fertilization on plant biomass production under elevated CO₂, we compared the response of N-fertilized tallgrass prairie at ambient and twice-ambient CO₂ levels over a 2-year period. Native tallgrass prairie plots (4.5 m diameter) were exposed continuously (24 h) to ambient and twice-ambient CO₂ from 1 April to 26 October. We compared our results to an unfertilized companion experiment on the same research site. Above- and belowground biomass production and leaf area of fertilized plots were greater with elevated than ambient CO₂ in both years. The increase in biomass at high CO₂ occurred mainly aboveground in 1991, a dry year, and belowground in 1990, a wet year. Nitrogen concentration was lower in plants exposed to elevated CO₂, but total standing crop N was greater at high CO₂. Increased root biomass under elevated CO₂ apparently increased N uptake. The biomass production response to elevated CO₂ was much greater on N-fertilized than unfertilized prairie, particularly in the dry year. We conclude that biomass production response to elevated CO₂ was suppressed by N limitation in years with below-normal precipitation. Reduced N concentration in above- and belowground biomass could slow microbial degradation of soil organic matter and surface litter, thereby exacerbating N limitation in the long term.     

The chemistry of wet deposition for a tallgrass prairie ecosystem: inputs and interactions with plant canopies

The chemistry and nutrient inputs of wet deposition, and the N chemistry of throughfall, were characterized for a tallgrass prairie in north-central Kansas. Dominant ions in wetfall were NH ₄ ⁺ , Ca²⁺, H⁺, NO ₃ ^- , and SO ₄ ^2- ; weighted mean pH was 4.79. Principal sources of ions appeared to be natural emissions and wind-blown soils. Concentrations of NO ₃ ^- -N, NH ₄ ⁺ -N, and organic N in wet deposition were 0.31, 0.30, and 0.17 mg/L, respectively, resulting in N inputs of 2.5, 2.5, and 1.4 kg · ha^-1 · yr^-1. Comparisons with bulk precipitation suggested that at least 50% of atmospheric N inputs were from dry deposition. Concentrations of NO ₃ ^- -N, NH ₄ ⁺ -N, and organic N in unburned prairie throughfall were 0.27, 0.28, and 1.28 mg/L, and in burned prairie throughfall were 0.33, 0.37, and 0.91 mg/L, respectively. The prairie canopy intercepted up to 48% of incident precipitation. Lower inorganic N and higher organic N concentrations in throughfall relative to wet deposition probably resulted from leaf uptake of N and immobilization by microbes associated with the standing dead plant materials of the prairie canopy. The removal of these materials by fire is important in maintaining N availability for tallgrass prairie. Much of the N immobilization appeared to have been of N that was supplied to the prairie canopy by dry deposition.