Steven S. Perakis

Global patterns of terrestrial biological nitrogen (N2) fixation in natural ecosystems

Human activities have clearly caused dramatic alterations of the terrestrial nitrogen cycle, and analyses of the extent and effects of such changes are now common in the scientific literature. However, any attempt to evaluate N cycling processes within ecosystems, as well as anthropogenic influences on the N cycle, requires an understanding of the magnitude of inputs via biological nitrogen fixation (BNF). Although there have been many studies addressing the microbiology, physiology, and magnitude of N fixation at local scales, there are very few estimates of BNF over large scales. We utilized >100 preexisting published estimates of BNF to generate biome- and global-level estimates of biological N fixation. We also used net primary productivity (NPP) and evapotranspiration (ET) estimates from the Century terrestrial ecosystem model to examine global relationships between these variables and BNF as well as to compare observed and Century-modeled BNF. Our data-based estimates showed a strong positive relationship between ecosystem ET and BNF, and our analyses suggest that while the models simple relationships for BNF predict broad scale patterns, they do not capture much of the variability or magnitude of published rates. Patterns of BNF were also similar to patterns of ecosystem NPP. Our “best estimate” of potential nitrogen fixation by natural ecosystems is ∼195 Tg N yr−1, with a range of 100–290 Tg N yr−1. Although these estimates do not account for the decrease in natural N fixation due to cultivation, this would not dramatically alter our estimate, as the greatest reductions in area have occurred in systems characterized by relatively low rates of N fixation (e.g., grasslands). Although our estimate of BNF in natural ecosystems is similar to previously published estimates of terrestrial BNF, we believe that this study provides a more documented, constrained estimate of this important flux.

Nitrogen loss from unpolluted South American forests mainly via dissolved organic compounds

Conceptual and numerical models of nitrogen cycling in temperate forests assume that nitrogen is lost from these ecosystems predominantly by way of inorganic forms, such as nitrate and ammonium ions. Of these, nitrate is thought to be particularly mobile, being responsible for nitrogen loss to deep soil and stream waters. But human activities—such as fossil fuel combustion, fertilizer production and land-use change—have substantially altered the nitrogen cycle over large regions, making it difficult to separate natural aspects of nitrogen cycling from those induced by human perturbations. Here we report stream chemistry data from 100 unpolluted primary forests in temperate South America. Although the sites exhibit a broad range of environmental factors that influence ecosystem nutrient cycles (such as climate, parent material, time of ecosystem development, topography and biotic diversity), we observed a remarkably consistent pattern of nitrogen loss across all forests. In contrast to findings from forests in polluted regions, streamwater nitrate concentrations are exceedingly low, such that nitrate to ammonium ratios were less than unity, and dissolved organic nitrogen is responsible for the majority of nitrogen losses from these forests. We therefore suggest that organic nitrogen losses should be considered in models of forest nutrient cycling, which could help to explain observations of nutrient limitation in temperate forest ecosystems.

Effects of nitrogen deposition and empirical nitrogen critical loads for ecoregions of the United States

Human activity in the last century has led to a significant increase in nitrogen (N) emissions and atmospheric deposition. This N deposition has reached a level that has caused or is likely to cause alterations to the structure and function of many ecosystems across the United States. One approach for quantifying the deposition of pollution that would be harmful to ecosystems is the determination of critical loads. A critical load is defined as the input of a pollutant below which no detrimental ecological effects occur over the long-term according to present knowledge. The objectives of this project were to synthesize current research relating atmospheric N deposition to effects on terrestrial and freshwater ecosystems in the United States, and to estimate associated empirical N critical loads. The receptors considered included freshwater diatoms, mycorrhizal fungi, lichens, bryophytes, herbaceous plants, shrubs, and trees. Ecosystem impacts included: (1) biogeochemical responses and (2) individual species, population, and community responses. Biogeochemical responses included increased N mineralization and nitrification (and N availability for plant and microbial uptake), increased gaseous N losses (ammonia volatilization, nitric and nitrous oxide from nitrification and denitrification), and increased N leaching. Individual species, population, and community responses included increased tissue N, physiological and nutrient imbalances, increased growth, altered root : shoot ratios, increased susceptibility to secondary stresses, altered fire regime, shifts in competitive interactions and community composition, changes in species richness and other measures of biodiversity, and increases in invasive species.

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.

NITROGEN RETENTION ACROSS A GRADIENT OF 15N ADDITIONS TO AN UNPOLLUTED TEMPERATE FOREST SOIL IN CHILE

Accelerated nitrogen (N) inputs can drive nonlinear changes in N cycling, retention, and loss in forest ecosystems. Nitrogen processing in soils is critical to understanding these changes, since soils typically are the largest N sink in forests. To elucidate soil mechanisms that underlie shifts in N cycling across a wide gradient of N supply, we added 15NH415NO3 at nine treatment levels ranging in geometric sequence from 0.2 kg to 640 kg N·ha−1·yr−1 to an unpolluted old-growth temperate forest in southern Chile. We recovered roughly half of 15N tracers in 0–25 cm of soil, primarily in the surface 10 cm. Low to moderate rates of N supply failed to stimulate N leaching, which suggests that most unrecovered 15N was transferred from soils to unmeasured sinks above ground. However, soil solution losses of nitrate increased sharply at inputs >160 kg N·ha−1·yr−1, corresponding to a threshold of elevated soil N availability and declining 15N retention in soil. Soil organic matter (<5.6 mm) dominated tracer retention at low rates of N input, but coarse roots and particulate organic matter became increasingly important at higher N supply. Coarse roots and particulate organic matter together accounted for 38% of recovered 15N in soils at the highest N inputs and may explain a substantial fraction of the “missing N” often reported in studies of fates of N inputs to forests. Contrary to expectations, N additions did not stimulate gross N cycling, potential nitrification, or ammonium oxidizer populations. Our results indicate that the nonlinearity in N retention and loss resulted directly from excessive N supply relative to sinks, independent of plant–soil–microbial feedbacks. However, N additions did induce a sharp decrease in microbial biomass C:N that is predicted by N saturation theory, and which could increase long-term N storage in soil organic matter by lowering the critical C:N ratio for net N mineralization. All measured sinks accumulated 15N tracers across the full gradient of N supply, suggesting that short-term nonlinearity in N retention resulted from saturation of uptake kinetics, not uptake capacity, in plant, soil, and microbial pools.

Nutrient Vectors and Riparian Processing: A Review with Special Reference to African Semiarid Savanna Ecosystems

A bstractThis review article describes vectors for nitrogen and phosphorus delivery to riparian zones in semiarid African savannas, the processing of nutrients in the riparian zone and the effect of disturbance on these processes. Semiarid savannas exhibit sharp seasonality, complex hillslope hydrology and high spatial heterogeneity, all of which ultimately impact nutrient fluxes between riparian, upland and aquatic environments. Our review shows that strong environmental drivers such as fire and herbivory enhance nitrogen, phosphorus and sediment transport to lower slope positions by shaping vegetative patterns. These vectors differ significantly from other arid and semiarid ecosystems, and from mesic ecosystems where the impact of fire and herbivory are less pronounced and less predictable. Also unique is the presence of sodic soils in certain hillslopes, which substantially alters hydrological flowpaths and may act as a trap where nitrogen is immobilized while sediment and phosphorus transport is enhanced. Nutrients and sediments are also deposited in the riparian zone during seasonal, intermittent floods while, during the dry season, subsurface movement of water from the stream into riparian soils and vegetation further enrich riparian zones with nutrients. As is found in mesic ecosystems, nutrients are immobilized in semiarid riparian corridors through microbial and plant uptake, whereas dissimilatory processes such as denitrification may be important where labile nitrogen and carbon are in adequate supply and physical conditions are suitable—such as in seeps, wallows created by animals, ephemeral wetlands and stream edges. Interaction between temporal hydrologic connectivity and spatial heterogeneity are disrupted by disturbances such as large floods and extended droughts, which may convert certain riparian patches from sinks to sources for nitrogen and phosphorus. In the face of increasing anthropogenic pressure, the scientific challenges are to provide a basic understanding of riparian biogeochemistry in semiarid African savannas to adequately address the temporal and spatial impact of disturbances, and to apply this knowledge to better regional land and water management. An integrated, multidisciplinary approach applied in protected as well as human-disturbed ecosystems in southern Africa is essential for underpinning a strong environmental basis for sustainable human-related expansion.

Biogeochemistry of a temperate forest nitrogen gradient

Wide natural gradients of soil nitrogen (N) can be used to examine fundamental relationships between plant-soil-microbial N cycling and hydrologic N loss, and to test N-saturation theory as a general framework for understanding ecosystem N dynamics. We characterized plant production, N uptake and return in litterfall, soil gross and net N mineralization rates, and hydrologic N losses of nine Douglas-fir (Pseudotsuga menziesii) forests across a wide soil N gradient in the Oregon Coast Range (U.S.A.). Surface mineral soil N (0-10 cm) ranged nearly three-fold from 0.29% to 0.78% N, and in contrast to predictions of N-saturation theory, was linearly related to 10-fold variation in net N mineralization, from 8 to 82 kg N.ha(-1) x yr(-1). Net N mineralization was unrelated to soil C:N, soil texture, precipitation, and temperature differences among sites. Net nitrification was negatively related to soil pH, and accounted for <20% of net N mineralization at low-N sites, increasing to 85-100% of net N mineralization at intermediate- and high-N sites. The ratio of net: gross N mineralization and nitrification increased along the gradient, indicating progressive saturation of microbial N demands at high soil N. Aboveground N uptake by plants increased asymptotically with net N mineralization to a peak of approximately 35 kg N.ha(-1) x yr(-1). Aboveground net primary production per unit net N mineralization varied inversely with soil N, suggesting progressive saturation of plant N demands at high soil N. Hydrologic N losses were dominated by dissolved organic N at low-N sites, with increased nitrate loss causing a shift to dominance by nitrate at high-N sites, particularly where net nitrification exceeded plant N demands. With the exception of N mineralization patterns, our results broadly support the application of the N-saturation model developed from studies of anthropogenic N deposition to understand N cycling and saturation of plant and microbial sinks along natural soil N gradients. This convergence of behavior in unpolluted and polluted forest N cycles suggests that where future reductions in deposition to polluted sites do occur, symptoms of N saturation are most likely to persist where soil N content remains elevated.

Terrestrial C sequestration at elevated CO2 and temperature : the role of dissolved organic N loss

We used a simple model of carbon-nitrogen (C-N) interactions in terrestrial ecosystems to examine the responses to elevated CO2 and to elevated CO2 plus warming in ecosystems that had the same total nitrogen loss but that differed in the ratio of dissolved organic nitrogen (DON) to dissolved inorganic nitrogen (DIN) loss. We postulate that DIN losses can be curtailed by higher N demand in response to elevated CO2, but that DON losses cannot. We also examined simulations in which DON losses were held constant, were proportional to the amount of soil organic matter, were proportional to the soil C:N ratio, or were proportional to the rate of decomposition. We found that the mode of N loss made little difference to the short-term (,60 years) rate of carbon sequestration by the ecosystem, but high DON losses resulted in much lower carbon sequestration in the long term than did low DON losses. In the short term, C sequestration was fueled by an internal redistribution of N from soils to vegetation and by increases in the C:N ratio of soils and vegetation. This sequestration was about three times larger with elevated CO 2 and warming than with elevated CO2 alone. After year 60, C sequestration was fueled by a net accu- mulation of N in the ecosystem, and the rate of sequestration was about the same with elevated CO2 and warming as with elevated CO2 alone. With high DON losses, the ecosystem either sequestered C slowly after year 60 (when DON losses were constant or proportional to soil organic matter) or lost C (when DON losses were proportional to the soil C:N ratio or to decomposition). We conclude that changes in long-term C sequestration depend not only on the magnitude of N losses, but also on the form of those losses.

Convergence of soil nitrogen isotopes across global climate gradients

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

Interannual variation of carbon fluxes from three contrasting evergreen forests: the role of forest dynamics and climate

Interannual variation of carbon fluxes can be attributed to a number of biotic and abiotic controls that operate at different spatial and temporal scales. Type and frequency of disturbance, forest dynamics, and climate regimes are important sources of variability. Assessing the variability of carbon fluxes from these specific sources can enhance the interpretation of past and current observations. Being able to separate the variability caused by forest dynamics from that induced by climate will also give us the ability to determine if the current observed carbon fluxes are within an expected range or whether the ecosystem is undergoing unexpected change. Sources of interannual variation in ecosystem carbon fluxes from three evergreen ecosystems, a tropical, a temperate coniferous, and a boreal forest, were explored using the simulation model STANDCARB. We identified key processes that introduced variation in annual fluxes, but their relative importance differed among the ecosystems studied. In the tropical site, intrinsic forest dynamics contributed approximately 30% of the total variation in annual carbon fluxes. In the temperate and boreal sites, where many forest processes occur over longer temporal scales than those at the tropical site, climate controlled more of the variation among annual fluxes. These results suggest that climate-related variability affects the rates of carbon exchange differently among sites. Simulations in which temperature, precipitation, and radiation varied from year to year (based on historical records of climate variation) had less net carbon stores than simulations in which these variables were held constant (based on historical records of monthly average climate), a result caused by the functional relationship between temperature and respiration. This suggests that, under a more variable temperature regime, large respiratory pulses may become more frequent and high enough to cause a reduction in ecosystem carbon stores. Our results also show that the variation of annual carbon fluxes poses an important challenge in our ability to determine whether an ecosystem is a source, a sink, or is neutral in regard to CO2 at longer timescales. In simulations where climate change negatively affected ecosystem carbon stores, there was a 20% chance of committing Type II error, even with 20 years of sequential data.