William B. Bowden

The biogeochemistry of nitrogen in freshwater wetlands

The biogeochemistry of N in freshwater wetlands is complicated by vegetation characteristics that range from annual herbs to perennial woodlands; by hydrologic characteristics that range from closed, precipitation-driven to tidal, riverine wetlands; and by the diversity of the nitrogen cycle itself. It is clear that sediments are the single largest pool of nitrogen in wetland ecosystems (100s to 1000s g N m-2) followed in rough order-of-magnitude decreases by plants and available inorganic nitrogen. Precipitation inputs (< 1–2 g N m-2 yr-1) are well known but other atmospheric inputs, e.g. dry deposition, are essentially unknown and could be as large or larger than wet deposition. Nitrogen fixation (acetylene reduction) is an important supplementary input in some wetlands (< < 1–3 g N m-2 yr-1) but is probably limited by the excess of fixed nitrogen usually present in wetland sediments.Plant uptake normally ranges from a few g N m-2 yr-1 to ∼ 35 g N m-2 yr-1 with extreme values of up to ∼ 100g N m-2 yr-1 Results of translocation experiments done to date may be misleading and may call for a reassessment of the magnitude of both plant uptake and leaching rates. Interactions between plant litter and decomposer microorganisms tend, over the short-term, to conserve nitrogen within the system in immobile forms. Later, decomposers release this nitrogen in forms and at rates that plants can efficiently reassimilate.The NO3 formed by nitrification (< 0.1 to 10 g N m-2 yr-1 has several fates which may tend to either conserve nitrogen (uptake and dissimilatory reduction to ammonium) or lead to its loss (denitrification). Both nitrification and denitrification operate at rates far below their potential and under proper conditions (e.g. draining or fluctuating water levels) may accelerate. However, virtually all estimates of denitrification rates in freshwater wetlands are based on measurements of potential denitrification, not actual denitrification and, as a consequence, the importance of denitrification in these ecosystems may have been greatly over estimated.In general, larger amounts of nitrogen cycle within freshwater wetlands than flow in or out. Except for closed, ombrotrophic systems this might seem an unusual characteristic for ecosystems that are dominated by the flux of water, however, two factors limit the opportunity for N loss. At any given time the fraction of nitrogen in wetlands that could be lost by hydrologic export is probably a small fraction of the potentially mineralizable nitrogen and is certainly a negligible fraction of the total nitrogen in the system. Second, in some cases freshwater wetlands may be hydrologically isolated so that the bulk of upland water flow may pass under (in the case of floating mats) or by (in the case of riparian systems) the biotically active components of the wetland. This may explain the rather limited range of N loading rates real wetlands can accept in comparison to, for example, percolation columns or engineered marshes.

N uptake as a function of concentration in streams

Detailed studies of stream N uptake were conducted in a prairie reach and gallery forest reach of Kings Creek on the Konza Prairie Biological Station. Nutrient uptake rates were measured with multiple short-term enrichments of NO3− and NH4+ at constant addition rates in the spring and summer of 1998. NH4+ uptake was also measured with 15N-NH4+ tracer additions and short-term unlabeled NH4+ additions at 12 stream sites across North America. Concurrent addition of a conservative tracer was used to account for dilution in all experiments. NH4+ uptake rate per unit area (Ut) was positively correlated to nutrient concentration across all sites (r2 = 0.41, log–log relationship). Relationships between concentration and Ut were used to determine whether the uptake was nonlinear (i.e., kinetic uptake primarily limited by the biotic capacity of microorganisms to accumulate nutrients) or linear (e.g., limited by mass transport into stream biofilms). In all systems, Ut was lower at ambient concentrations than at elevated concentrations. Extrapolation from uptake measured from a series of increasing enrichments could be used to estimate ambient Ut. Linear extrapolation of Ut assuming the relationship passes through the origin and rates measured at 1 elevated nutrient concentration underestimated ambient Ut by ∼3-fold. Uptake rates were saturated under some but not all conditions of enrichment; in some cases there was no saturation up to 50 μmol/L. The absolute concentration at which Ut was saturated in Kings Creek varied among reaches and nutrients. Uptake rates of NH4+ at ambient concentrations in all streams were higher than would be expected, assuming Ut does not saturate with increasing concentrations. At ambient nutrient concentrations in unpolluted streams, Ut is probably limited to some degree by the kinetic uptake capacity of stream biota. Mass transfer velocity from the water column is generally greater than would be expected given typical diffusion rates, underscoring the importance of advective transport. Given the short-term spikes in nutrient concentrations that can occur in streams (e.g., in response to storm events), Ut may not saturate, even at high concentrations.

Can uptake length in streams be determined by nutrient addition experiments? Results from an interbiome comparison study

Nutrient uptake length is an important parameter for quantifying nutrient cycling in streams. Although nutrient tracer additions are the preferred method for measuring uptake length under ambient nutrient concentrations, short-term nutrient addition experiments have more frequently been used to estimate uptake length in streams. Theoretical analysis of the relationship between uptake length determined by nutrient addition experiments (SW′) and uptake length determined by tracer additions (SW) predicted that SW′ should be consistently longer than SW, and that the overestimate of uptake length by SW′ should be related to the level of nutrient addition above ambient concentrations and the degree of nutrient limitation. To test these predictions, we used data from an interbiome study of NH4+ uptake length in which 15NH4+ tracer and short-term NH4+ addition experiments were performed in 10 streams using a uniform experimental approach. The experimental results largely confirmed the theoretical predictions: SW′ was consistently longer than SW and SW′:SW ratios were directly related to the level of NH4+ addition and to indicators of N limitation. The experimentally derived SW′:SW ratios were used with the theoretical results to infer the N limitation status of each stream. Together, the theoretical and experimental results showed that tracer experiments should be used whenever possible to determine nutrient uptake length in streams. Nutrient addition experiments may be useful for comparing uptake lengths between different streams or different times in the same stream, however, provided that nutrient additions are kept as low as possible and of similar magnitude.

Gaseous nitrogen emmissions from undisturbed terrestrial ecosystems: An assessment of their impacts on local and global nitrogen budgets

There is increasing interest in the importance of nitrogen gas emissions from natural (non-agricultural) ecosystems with respect to local as well as global nitrogen budgets and with respect to the effects of nitrogen oxides on atmospheric ozone levels and global warming. The volatile forms of nitrogen of common interest are ammonia (NH3), nitrous oxide, (N2O), dinitrogen (N2), and NOx (principally NO + NO2). It is often difficult to attribute emissions of these compounds from soils to a single process because they are produced by a variety of common biogeochemical mechanisms. Although environmental conditions in the soil often appear to favor nitrogen gas emissions, the potential nitrogen gas emission rate from undisturbed ecosystems is rarely approached. The best estimates to date suggest that nitrogen gas emission rates from undisturbed ecosystems typically range from > 1 to perhaps 10 or 20 kg N ha-1 yr-1. Under certain conditions, however, emission rates may be much higher. For example, excreta from animals in grasslands may elevate ammonia volatilization up to 100 kg N ha-1 yr-1 depending on grazer density; tidal input of nutrients to coastal wetlands may support denitrification rates of several hundred kg N ha-1 yr-1 . Excepting such cases, gaseous nitrogen losses are probably a small component of the local nitrogen budget in most undisturbed ecosystems. However, emissions from undisturbed soils are an important component of the global source strengths for (N2O + N2), N2O and NOx (50%, 21%, and 10% respectively). Emission rates of N2O from natural ecosystems are higher than assumed previously by perhaps 10 times. Large-scale disturbance may have a stimulatory effect on nitrogen emission rates which could have important effects on global nitrogen budgets. There is a need for more sophisticated methods to account for natural temporal and spatial variations of emissions rates, to more accurately and precisely assess their global source strengths.

A Cross-System Comparison of Bacterial and Fungal Biomass in Detritus Pools of Headwater Streams

The absolute amount of microbial biomass and relative contribution of fungi and bacteria are expected to vary among types of organic matter (OM) within a stream and will vary among streams because of differences in organic matter quality and quantity. Common types of benthic detritus [leaves, small wood, and fine benthic organic matter (FBOM)] were sampled in 9 small (1st-3rd order) streams selected to represent a range of important controlling factors such as surrounding vegetation, detritus standing stocks, and water chemistry. Direct counts of bacteria and measurements of ergosterol (a fungal sterol) were used to describe variation in bacterial and fungal biomass. There were significant differences in bacterial abundance among types of organic matter with higher densities per unit mass of organic matter on fine particles relative to either leaves or wood surfaces. In contrast, ergosterol concentrations were significantly greater on leaves and wood, confirming the predominance of fungal biomass in these larger size classes. In general, bacterial abundance per unit organic matter was less variable than fungal biomass, suggesting bacteria will be a more predictable component of stream microbial communities. For 7 of the 9 streams, the standing stock of fine benthic organic matter was large enough that habitat-weighted reach-scale bacterial biomass was equal to or greater than fungal biomass. The quantities of leaves and small wood varied among streams such that the relative contribution of reach-scale fungal biomass ranged from 10% to as much as 90% of microbial biomass. Ergosterol concentrations were positively associated with substrate C:N ratio while bacterial abundance was negatively correlated with C:N. Both these relationships are confounded by particle size, i.e., leaves and wood had higher C:N than fine benthic organic matter. There was a weak positive relationship between bacterial abundance and streamwater soluble reactive phosphorus concentration, but no apparent pattern between either bacteria or fungi and streamwater dissolved inorganic nitrogen. The variation in microbial biomass per unit organic matter and the relative abundance of different types of organic matter contributed equally to driving differences in total microbial biomass at the reach scale.

LONG‐TERM RESPONSES OF THE KUPARUK RIVER ECOSYSTEM TO PHOSPHORUS FERTILIZATION

A long-term stream fertilization experiment was performed to evaluate the potential eutrophication of an arctic stream ecosystem. During 16 years of summer phosphorus (H3PO4) fertilization, we observed a dramatic change in the community structure of the Kuparuk River on the North Slope of Alaska. A positive response to fertilization was observed at all trophic levels with increases in epilithic algal stocks, some insect densities, and fish growth rates. After approximately eight years of P fertilization, bryophytes (mosses) replaced epilithic diatoms as the dominant primary producers in the Kuparuk River. The moss impacted NH4+ uptake rates, benthic gross primary production, habitat structure, and insect abundance and species composition. This study documents the long-term changes in an arctic tundra stream in response to nutrient enrichment. Predicting stream ecosystem responses to chronic perturbation requires long-term observation and experiments.

Riparian nitrogen dynamics in two geomorphologically distinct tropical rain forest watersheds: subsurface solute patterns

Nitrate, ammonium, dissolved organic N, and dissolved oxygen were measured in stream water and shallow groundwater in the riparian zones of two tropical watersheds with different soils and geomorphology. At both sites, concentrations of dissolved inorganic N (DIN; NH4+- and NO3−-N) were low in stream water (< 110 ug/L). Markedly different patterns in DIN were observed in groundwater collected at the two sites. At the first site (Icacos watershed), DIN in upslope groundwater was dominated by NO3−-N (550 ug/L) and oxygen concentrations were high (5.2 mg/L). As groundwater moved through the floodplain and to the stream, DIN shifted to dominance by NH4+-N (200–700 ug/L) and groundwater was often anoxic. At the second site (Bisley watershed), average concentrations of total dissolved nitrogen were considerably lower (300 ug/L) than at Icacos (600 ug/L), and the dominant form of nitrogen was DON rather than inorganic N. Concentrations of NH4+ and NO3− were similar throughout the riparian zone at Bisley, but concentrations of DON declined from upslope wells to stream water.Differences in speciation and concentration of nitrogen in groundwater collected at the two sites appear to be controlled by differences in redox conditions and accessibility of dissolved N to plant roots, which are themselves the result of geomorphological differences between the two watersheds. At the Icacos site, a deep layer of coarse sand conducts subsurface water to the stream below the rooting zone of riparian vegetation and through zones of strong horizontal redox zonation. At the Bisley site, infiltration is impeded by dense clays and saturated flow passes through the variably oxidized rooting zone. At both sites, hydrologic export of nitrogen is controlled by intense biotic activity in the riparian zone. However, geomorphology appears to strongly modify the importance of specific biotic components.

Rapid N^2 Fixation in Pines, Alder, and Locust: Evidence From the Sandbox Ecosystems Study

Not all nitrogen (N) inputs have been accounted for in forested ecosystems. We sought to account for N2 fixation and dry deposition using a lysimeter mass—balance approach. Large sand—filled, field lysimeters were used to construct 5—yr nitrogen budgets for two N2—fixing trees, two pines, and a nonvegetated control soil. This approach is a promising and straightforward technique for quantifying otherwise difficult—to—measure fluxes. Accurate assessment of changes in N storage combined with direct measurement of N inputs in precipitation and losses from leaching allowed as to estimate fluxes. Gains of N in pine systems were greatest in vegetation and litter, overshadowing combined losses from mineral soil and leaching by about threefold. Rapid acetylene reduction in pine rhizospheres and in cultures from washed roots suggests that unexplained gains are due to associative N2 fixation. These results provide strong evidence for N2 fixation in pine systems of °50 kg°ha—1°yr—1 N. The symbiotic N2—fixing trees black locust and black alder fixed 2 and 5 times more N2, respectively, than did pines. In all systems, input in precipitation and dry deposition were relatively unimportant to the N budget. Unexplained losses of N from the nonvegetated control suggest that denitrification is an important flux. Mineral soil organic matter declined sharply and significantly in pines (20%) and even more so in the nonvegetated control (40%). Symbiotic N2—fixing trees caused a small, nonsignificant increase in mineral soil organic matter and large, significant increases in litter layer organic matter and large, significant increases in litter layer organic matter. Bulk density (0—20 cm) declined by 5% under symbiotic N2—fixing trees and increased by 5% in one pine sandbox. Correction for soil expansion or collapse did not greatly alter estimates of unexplained N or N2 fixation. Pines with rhizospheres that fix N2 at the rates we observed might be used to restore degraded land and to create silvicultural systems that are N self sufficient. We first need to better understand the microbiology, tree genetics, and soil conditions that lead to rapid N2 fixation in pine ecosystems.

Carbon and nitrogen stoichiometry and nitrogen cycling rates in streams

Stoichiometric analyses can be used to investigate the linkages between N and C cycles and how these linkages influence biogeochemistry at many scales, from components of individual ecosystems up to the biosphere. N-specific NH4+ uptake rates were measured in eight streams using short-term 15N tracer additions, and C to N ratios (C:N) were determined from living and non-living organic matter collected from ten streams. These data were also compared to previously published data compiled from studies of lakes, ponds, wetlands, forests, and tundra. There was a significant negative relationship between C:N and N-specific uptake rate; C:N could account for 41% of the variance in N-specific uptake rate across all streams, and the relationship held in five of eight streams. Most of the variation in N-specific uptake rate was contributed by detrital and primary producer compartments with large values of C:N and small values for N-specific uptake rate. In streams, particulate materials are not as likely to move downstream as dissolved N, so if N is cycling in a particulate compartment, N retention is likely to be greater. Together, these data suggest that N retention may depend in part on C:N of living and non-living organic matter in streams. Factors that alter C:N of stream ecosystem compartments, such as removal of riparian vegetation or N fertilization, may influence the amount of retention attributed to these ecosystem compartments by causing shifts in stoichiometry. Our analysis suggests that C:N of ecosystem compartments can be used to link N-cycling models across streams.

Temporal variation of N and P uptake in 2 New Zealand streams

Abstract Temporal variation of nutrient uptake in streams may be large because nutrient uptake is driven by many factors that vary substantially over time. Although many studies have compared nutrient uptake among streams, the range and variation of nutrient uptake within streams is known only for a few streams and a few nutrients. We investigated the monthly variation of NH4+, NO3−, and PO43− uptake in 2 New Zealand streams over 1 y. To measure uptake, each nutrient was added individually along with a conservative tracer (Cl−) into each stream on 3 successive days in each month. Ambient nutrient concentrations were low and nutrients were efficiently removed from the water column, with maximum uptake velocities (vf) of 71, 12, and 11 mm/min for NH4+, NO3−, and PO43−, respectively. Nutrient uptake varied considerably during the year (CV = 37–109%), with shortest nutrient uptake lengths (Sw) and highest vf generally in spring and summer months. The range of vf occurring within the streams spanned 25 to 89% of the range of vf among other streams. The range of uptake rates (U) within the streams was lower, accounting for 2 to 40% of the range among other streams. Variation in Sw was largely explained by changes in velocity and effective depth. Physical factors (temperature, transient storage) and chlorophyll a were generally poor predictors of vf and U. There was little correlation in uptake among nutrients, suggesting different factors were responsible for uptake of each nutrient. Our results show that the range and variation of nutrient uptake within some streams can be large. Within-stream variation should be considered when comparing among streams and may be useful for understanding what factors drive nutrient uptake in streams.