Nitrogen saturation in stream ecosystems

The concept of nitrogen (N) saturation has organized the assessment of N loading in terrestrial ecosystems. Here we extend the concept to lotic ecosystems by coupling Michaelis-Menten kinetics and nutrient spiraling. We propose a series of saturation response types, which may be used to characterize the proximity of streams to N saturation. We conducted a series of short-term N releases using a tracer (15NO3-N) to measure uptake. Experiments were conducted in streams spanning a gradient of background N concentration. Uptake increased in four of six streams as NO3-N was incrementally elevated, indicating that these streams were not saturated. Uptake generally corresponded to Michaelis-Menten kinetics but deviated from the model in two streams where some other growth-critical factor may have been limiting. Proximity to saturation was correlated to background N concentration but was better predicted by the ratio of dissolved inorganic N (DIN) to soluble reactive phosphorus (SRP), suggesting phosphorus limitation in several high-N streams. Uptake velocity, a reflection of uptake efficiency, declined nonlinearly with increasing N amendment in all streams. At the same time, uptake velocity was highest in the low-N streams. Our conceptual model of N transport, uptake, and uptake efficiency suggests that, while streams may be active sites of N uptake on the landscape, N saturation contributes to nonlinear changes in stream N dynamics that correspond to decreased uptake efficiency.     

Urbanization and nutrient retention in freshwater riparian wetlands.

Urbanization can degrade water quality and alter watershed hydrology, with profound effects on the structure and function of both riparian wetlands (RWs) and aquatic ecosystems downstream. We used freshwater RWs in Fairfax County, Virginia, USA, as a model system to examine: (1) the effects of increasing urbanization (indexed by the percentage of impervious surface cover [%ISC] in the surrounding watershed) on nitrogen (N) and phosphorus (P) concentrations in surface soils and plant tissues, soil P saturation, and soil iron (Fe) chemistry; and (2) relationships between RW soil and plant nutrient chemistries vs. the physical and biotic integrity of adjacent streams. Soil total P and NaOH-extractable P (representing P bound to aluminum [Al] and Fe hydrous oxides) varied significantly but nonlinearly with % ISC (r2 = 0.69 and 0.57, respectively); a similar pattern was found for soil P saturation but not for soil total N. Relationships were best described by second-order polynomial equations. Riparian wetlands appear to receive greater P loads in moderately (8.6-13.3% ISC) than in highly (25.1-29.1% ISC) urbanized watersheds. These observations are consistent with alterations in watershed hydrology that occur with increasing urbanization, directing water and nutrient flows away from natural RWs. Significant increases in total and crystalline soil Fe (r2 = 0.57 and 0.53, respectively) and decreases in relative soil Fe crystallinity with increasing %ISC suggest the mobilization and deposition of terrestrial sediments in RWs, likely due to construction activities in the surrounding watershed. Increases in RW plant tissue nutrient concentrations and %ISC in the surrounding watershed were negatively correlated with standard indices of the physical and biotic integrity of adjacent streams. In combination, these data suggest that nutrient and sediment inputs associated with urbanization and storm-water management are important variables that affect wetland ecosystem services, such as water quality improvement, in urbanizing landscapes.     

Land use change and nitrogen enrichment of a Rocky Mountain watershed.

Headwater ecosystems may have a limited threshold for retaining and removing nutrients delivered by certain types of land use. Nitrogen enrichment was studied in a Rocky Mountain watershed undergoing rapid expansion of population and residential development. Study sites were located along a 30-km transect from the headwaters of the Blue River to Lake Dillon, a major source of drinking water for Denver, Colorado. Ground water in residential areas with septic systems showed high concentrations of nitrate-N (4.96 +/- 1.22 mg/L, mean +/- SE), and approximately 40% of wells contained nitrate with delta15N values in the range of wastewater. Concentrations of dissolved inorganic nitrogen (DIN) in tributaries with residential development peaked during spring snowmelt as concentrations of DIN declined to below detection limits in undeveloped tributaries. Annual export of dissolved organic nitrogen (DON) was considerably lower in residential streams, suggesting a change in forms of N with development. The seasonal delta15N of algae in residential streams was intermediate between baseline values from undeveloped streams and stream algae grown on wastewater. Between 19% and 23% of the annual N export from developed tributaries was derived from septic systems, as estimated from the delta15N of algae. This range was similar to the amount of N export above background determined independently from mass-balance estimates. From a watershed perspective, total loading of N to the Blue River catchment from septic and municipal wastewater (2 kg x ha(-1) x yr(-1)) is currently less than the amount from background atmospheric sources (3 kg x ha(-1) x yr(-1)). Nonetheless, nitrate-N concentrations exceeded limits for safe drinking water in some groundwater wells (10 mg/L), residential streams showed elevated seasonal patterns of nitrate-N concentration and ratios of DIN to total dissolved phosphorus, and seasonal minimum concentrations of nitrate-N in Lake Dillon have increased exponentially to 80 microg/L over the last decade from an initial value near zero. Results suggest that isotopic ratios in autotrophs can be used to detect and quantify increases in N enrichment associated with land use change. The biotic capacity of headwater ecosystems to assimilate increases in inorganic N from residential development may be insufficient to prevent nitrogen enrichment over considerable distances and multiple aquatic ecosystems downstream.     

Changes in the dissolved nitrogen pool across land cover gradients in Wisconsin streams

Increases in anthropogenic nitrogen fixation have resulted in wide-scale enrichment of aquatic ecosystems. Existing biogeochemical theory suggests that N enrichment is associated with increasing concentrations of nitrate; however, dissolved organic nitrogen (DON) is often a major component of the total dissolved nitrogen (TDN) pool in streams and rivers, and its concentration can be significantly elevated in human-influenced basins. We examined N concentrations during summer base flow conditions in 324 Wisconsin streams to determine whether DON was a significant component of TDN and how its relative contribution changed across a gradient of increasing human (agriculture and urban) land use for 84 of these sites. Total dissolved nitrogen varied from 0.09 to 20.74 mg/L, and although DON was significantly higher in human-dominated basins relative to forested and mixed-cover basins, its concentration increased relatively slowly in response to increasing human land cover. This limited response reflected a replacement of wetland-derived DON in low-N streams by anthropogenic sources in human-dominated sites, such that net changes in DON were small across the land use gradient. Nitrate-N increased exponentially in response to greater human land cover, and NH4-N and NO2-N were present at low levels. Nitrite-N exceeded NH4-N at 20% of sites and reached a maximum concentration of 0.10 mg/L. This examination suggests that basic mechanisms driving N losses from old-growth forests subject to N saturation also shape the summertime N pool in Wisconsin streams, in addition to other processes dictated by landscape context. The overwhelming role of human land use in determining the relative and absolute composition of the summertime N pool included (1) rapid increases in NO3-N, (2) limited changes in DON, and (3) the unexpected occurrence of NO2-N. High (>3 mg/L) TDN conditions dominated by NO3-N, regardless of landscape context or forms of N inputs, indicate a state of "N hypersaturation", which appears to be increasingly common in human-influenced streams and rivers. Many sites in agriculturally rich areas had NO2-N and NO3-N concentrations that, if sustained, are at chronically toxic levels for sensitive aquatic biota, suggesting that N enrichment now has local consequences for resident stream biota in addition to contributing to coastal eutrophication.     

A reevaluation of the cultural eutrophication of Lake Okeechobee using multiproxy sediment records.

Lake Okeechobee, the hydrological lynchpin of the Everglades ecosystem, is the subject of an ambitious, multiagency restoration effort aimed at reducing phosphorus inputs and resulting algal blooms and impaired water clarity. This restoration is predicated on returning the lake to something closer to its predisturbance condition, but that goal has been challenged on the premise that the lake has always been eutrophic. The resolution of this debate and the appropriateness of the nutrient reduction goals thus depend on obtaining a reliable sediment record of past limnological conditions--the aim of this study. Because of the potential for severe sediment mixing from tropical storms, this investigation used multiple dating tools to examine the integrity of the sediment record and then analyzed proxies for nutrient enrichment, phytoplankton composition, and paleoproductivity. Sediment profiles for atmospheric pollutants, fertilizer contaminants, and radiocesium from three widely spaced cores showed good preservation of stratigraphic detail and coherence with the 210Pb chronologies. These results demonstrated that sediment stratigraphy is largely intact and retains a reliable record of limnological change. Geochemical proxies provide strong evidence of increased nutrient loading beginning ca. 1950. Concentrations of sediment P double, and N:P and C:N ratios drop, while those for N isotopes (delta15N) increase. At the same time, tracers of phosphate fertilizers (uranium, vanadium, and arsenic) rise. These changes are synchronous among cores and constitute a robust, internally consistent record of increasing water-column P. Biotic responses are manifested in higher concentrations and in changing composition of fossil algal pigments, including (1) large increases in the concentrations of chemically robust carotenoids, (2) corresponding decreases in the ratios of pigments from diatoms to chlorophyte and cyanobacterial algae, and (3) increases in UVR-photo-protective compounds indicating greater prevalence of surface algal blooms. This study provides strong evidence that Lake Okeechobee has experienced accelerated eutrophication linked with post-1950s land use changes in its watershed, a conclusion consistent with the nutrient reduction goals of the Lake Okeechobee Protection Program. The results contradict recent claims that the lake's trophic state has not changed over time, as well as the assertion that sediments of large shallow lakes cannot support a reliable chronology of past events.     

Fish distributions and nutrient cycling in streams: can fish create biogeochemical hotspots?

Rates of biogeochemical processes often vary widely in space and time, and characterizing this variation is critical for understanding ecosystem functioning. In streams, spatial hotspots of nutrient transformations are generally attributed to physical and microbial processes. Here we examine the potential for heterogeneous distributions of fish to generate hotspots of nutrient recycling. We measured nitrogen (N) and phosphorus (P) excretion rates of 47 species of fish in an N-limited Neotropical stream, and we combined these data with population densities in each of 49 stream channel units to estimate unit- and reach-scale nutrient recycling. Species varied widely in rates of N and P excretion as well as excreted N:P ratios (6-176 molar). At the reach scale, fish excretion could meet >75% of ecosystem demand for dissolved inorganic N and turn over the ambient NH4 pool in <0.3 km. Areal N excretion estimates varied 47-fold among channel units, suggesting that fish distributions could influence local N availability. P excretion rates varied 14-fold among units but were low relative to ambient concentrations. Spatial variation in aggregate nutrient excretion by fish reflected the effects of habitat characteristics (depth, water velocity) on community structure (body size, density, species composition), and the preference of large-bodied species for deep runs was particularly important. We conclude that the spatial distribution of fish could indeed create hotspots of nutrient recycling during the dry season in this species-rich tropical stream. The prevalence of patchy distributions of stream fish and invertebrates suggests that hotspots of consumer nutrient recycling may often occur in stream ecosystems.     

Assessing stream restoration effectiveness at reducing nitrogen export to downstream waters

The degradation of headwater streams is common in urbanized coastal areas, and the role these streams play in contributing to downstream pollution is a concern among natural resource managers and policy makers. Thus, many urban stream restoration efforts are increasingly focused on reducing the downstream flux of pollutants. In regions that suffer from coastal eutrophication, it is unclear whether stream restoration does in fact reduce nitrogen (N) flux to downstream waters and, if so, by how much and at what cost. In this paper, we evaluate whether stream restoration implemented to improve water quality of urban and suburban streams in the Chesapeake Bay region, USA, is effective at reducing the export of N in stream flow to downstream waters. We assessed the effectiveness of restored streams positioned in the upland vs. lowland regions of Coastal Plain watershed during both average and stormflow conditions. We found that, during periods of low discharge, lowland streams that receive minor N inputs from groundwater or bank seepage reduced in-stream N fluxes. Furthermore, lowland streams with the highest N concentrations and lowest discharge were the most effective. During periods of high flow, only those restoration projects that converted lowland streams to stream-wetland complexes seemed to be effective at reducing N fluxes, presumably because the design promoted the spillover of stream flow onto adjacent floodplains and wetlands. The observed N-removal rates were relatively high for stream ecosystems, and on the order of 5% of the inputs to the watershed. The dominant forms of N entering restored reaches varied during low and high flows, indicating that N uptake and retention were controlled by distinctive processes during different hydrological conditions. Therefore, in order for stream restoration to effectively reduce N fluxes exported to downstream waters, restoration design should include features that enhance the processing and retention of different forms of N, and for a wide range of flow conditions. The use of strategic designs that match the dominant attributes of a stream such as position in the watershed, influence of groundwater, dominant flow conditions, and N concentrations is crucial to assure the success of restoration.     

Examining the coupling of carbon and nitrogen cycles in Appalachian streams: the role of dissolved organic nitrogen

Although regional and global models of nitrogen (N) cycling typically focus on nitrate, dissolved organic nitrogen (DON) is the dominant form of nitrogen export from many watersheds and thus the dominant form of dissolved N in many streams. Our understanding of the processes controlling DON export from temperate forests is poor. In pristine systems, where biological N limitation is common, N contained in recalcitrant organic matter (OM) can dominate watershed N losses. This recalcitrant OM often has moderately constrained carbon:nitrogen (C:N) molar ratios (approximately 25-55) and therefore, greater DON losses should be observed in sites where there is greater total dissolved organic carbon (DOC) loss. In regions where anthropogenic N pollution is high, it has been suggested that increased inorganic N availability can reduce biological demand for organic N and therefore increase watershed DON losses. This would result in a positive correlation between inorganic and organic N concentrations across sites with varying N availability. In four repeated synoptic surveys of stream water chemistry from forested watersheds along an N loading gradient in the southern Appalachians, we found surprisingly little correlation between DON and DOC concentrations. Further, we found that DON concentrations were always significantly correlated with watershed N loading and stream water [NO3-] but that the direction of this relationship was negative in three of the four surveys. The C:N molar ratio of dissolved organic matter (DOM) in streams draining watersheds with high N deposition was very high relative to other freshwaters. This finding, together with results from bioavailability assays in which we directly manipulated C and N availabilities, suggests that heterotrophic demand for labile C can increase as a result of dissolved inorganic N (DIN) loading, and that heterotrophs can preferentially remove N-rich molecules from DOM. These results are inconsistent with the two prevailing hypotheses that dominate interpretations of watershed DON loss. Therefore, we propose a new hypothesis, the indirect carbon control hypothesis, which recognizes that heterotrophic demand for N-rich DOM can keep stream water DON concentrations low when N is not limiting and heterotrophic demand for labile C is high.     

Grazer traits, competition, and carbon sources to a headwater-stream food web

We investigated the effect of grazing by a dominant invertebrate grazer (the caddisfly Glossosoma penitum) on the energy sources used by other consumers in a headwater-stream food web. Stable isotope studies in small, forested streams in northern California have shown that G. penitum larvae derive most of their carbon from algae, despite low algal standing crops. We hypothesized that the caddisfly competes with other primary consumers (including mayflies) for algal food and increases their reliance on terrestrial detritus. Because Glossosoma are abundant and defended from predators by stone cases, their consumption of algal energy may reduce its transfer up the food chain. We removed Glossosoma (natural densities >1000 caddisflies/m2) from five approximately 4 m2) stream sections during the summer of 2000 and measured responses of algae, invertebrate primary consumers, and invertebrate predators. The treatment reduced Glossosoma biomass by 80-90%. We observed a doubling in chlorophyll a per area in sections with reduced Glossosoma abundance and aggregative increases in the biomass of undefended primary consumers. Heptageniid mayfly larvae consumed more algae (as measured by stable carbon isotope ratios and gut content analysis) in caddisfly removal plots at the end of the 60-day experiment, although not after one month. We did not see isotopic evidence of increased algal carbon in invertebrate predators, however. Patterns of caddisfly and mayfly diets in the surrounding watershed suggested that mayfly diets are variable and include algae and detrital carbon in variable proportions, but scraping caddisflies consume primarily algae. Caddisfly and mayfly diets are more similar in larger, more productive streams where the mayflies assimilate more algae. Isotopic analysis, in combination with measurements of macroinvertebrate abundance and biomass in unmanipulated plots, suggested that a substantial portion of the invertebrate community (>50% of biomass) was supported, at least partially, by local algal carbon during midsummer. These data suggest that algae may be more important to community dynamics in headwater streams than their relatively low productivity would suggest. Through their high densities and relative invulnerability to predation, armored grazers may also affect community structure and flow of algal and detrital carbon in headwater streams.     

Environmental controls on the landscape-scale biogeography of stream bacterial communities

We determined the biogeographical distributions of stream bacteria and the biogeochemical factors that best explained heterogeneity for 23 locations within the Hubbard Brook watershed, a 3000-ha forested watershed in New Hampshire, USA. Our goal was to assess the factor, or set of factors, responsible for generating the biogeographical patterns exhibited by microorganisms at the landscape scale. We used DNA fingerprinting to characterize bacteria inhabiting fine benthic organic matter (FBOM) because of their important influence on stream nutrient dynamics. Across the watershed, streams of similar pH had similar FBOM bacterial communities. Streamwater pH was the single variable most strongly correlated with the relative distance between communities (Spearman's p = 0.66, P < 0.001) although there were other contributing factors, including the quality of the fine benthic organic matter and the amount of dissolved organic carbon and nitrogen in the stream water (P < 0.05 for each). There was no evidence of an effect of geographic distance on bacterial community composition, suggesting that dispersal limitation has little influence on the observed biogeographical patterns in streams across this landscape. Cloning and sequencing of small-subunit rRNA genes confirmed the DNA fingerprinting results and revealed strong shifts among bacterial groups along the pH gradient. With an increase in streamwater pH, the abundance of acidobacteria in the FBOM bacterial community decreased (from 71% to 38%), and the abundance of proteobacteria increased (from 11% to 47%). Together these results suggest that microorganisms, like "macro"-organisms, do exhibit biogeographical patterns at the landscape scale and that these patterns may be predictable based on biogeochemical factors.     

15N-Nitrate signature in low-order streams: effects of land cover and agricultural practices.

Many studies have shown that intensive agricultural practices significantly increase the nitrogen concentration of stream surface waters, but it remains difficult to identify, quantify, and differentiate between terrestrial and in-stream sources or sinks of nitrogen, and rates of transformation. In this study we used the delta15N-NO3 signature in a watershed dominated by agriculture as an integrating marker to trace (1) the effects of the land cover and agricultural practices on stream-water N concentration in the upstream area of the hydrographic network, (2) influence of the in-stream processes on the NO3-N loads at the reach scale (100 m and 1000 m long), and (3) changes in delta15N-NO3 signature with increasing stream order (from first to third order). This study suggests that land cover and fertilization practices were the major determinants of delta15N-NO3 signature in first-order streams. NO3-N loads and delta15N-NO3 signature increased with fertilization intensity. Small changes in delta15N-NO3 signature and minor inputs of groundwater were observed along both types of reaches, suggesting the NO3-N load was slightly influenced by in-stream processes. The variability of NO3-N concentrations and delta15N signature decreased with increasing stream order, and the delta15N signature was positively correlated with watershed areas devoted to crops, supporting a dominant effect of agriculture compared to the effect of in-stream N processing. Consequently, land cover and fertilization practices are integrated in the natural isotopic signal at the third-order stream scale. The GIS analysis of the land cover coupled with natural-abundance isotope signature (delta15N) represents a potential tool to evaluate the effects of agricultural practices in rural catchments and the consequences of future changes in management policies at the regional scale.     

Watershed land use effects on lake water quality in Denmark

Mitigating nutrient losses from anthropogenic nonpoint sources is today of particular importance for improving the water quality of numerous freshwater lakes worldwide. Several empirical relationships between land use and in-lake water quality variables have been developed, but they are often weak, which can in part be attributed to lack of detailed information about land use activities or point sources. We examined a comprehensive data set comprising land use data, point-source information, and in-lake water quality for 414 Danish lakes. By excluding point-source-influenced lakes (n = 210), the strength in relationship (R2) between in-lake total nitrogen (TN) and total phosphorus (TP) concentrations and the proportion of agricultural land use in the watershed increased markedly, from 10-12% to 39-42% for deep lakes and from 10-12% to 21-23% for shallow lakes, with the highest increase for TN. Relationships between TP and agricultural land use were even stronger for lakes with rivers in their watershed (55%) compared to lakes without (28%), indicating that rivers mediate a stronger linkage between landscape activity and lake water quality by providing a "delivery" mechanism for excess nutrients in the watershed. When examining the effect of different near-freshwater land zones in contrast to the entire watershed, relationships generally improved with size of zone (25, 50, 100, 200, and 400 m from the edge of lake and streams) but were by far strongest using the entire watershed. The proportion of agricultural land use in the entire watershed was best in explaining lake water quality, both relative to estimated nutrient surplus at agricultural field level and near-lake land use, which somewhat contrasts typical strategies of management policies that mainly target agricultural nutrient applications and implementation of near-water buffer zones. This study suggests that transport mechanisms within the whole catchment are important for the nutrient export to lakes. Hence, the whole watershed should be considered when managing nutrient loadings to lakes, and future policies should ideally target measures that reduce the proportion of cultivated land in the watershed to successfully improve lake water quality.     

Are rivers just big streams? A pulse method to quantify nitrogen demand in a large river

Given recent focus on large rivers as conduits for excess nutrients to coastal zones, their role in processing and retaining nutrients has been overlooked and understudied. Empirical measurements of nutrient uptake in large rivers are lacking, despite a substantial body of knowledge on nutrient transport and removal in smaller streams. Researchers interested in nutrient transport by rivers (discharge >10000 L/s) are left to extrapolate riverine nutrient demand using a modeling framework or a mass balance approach. To begin to fill this knowledge gap, we present data using a pulse method to measure inorganic nitrogen. (N) transport and removal in the Upper Snake River, Wyoming, USA (seventh order, discharge 12000 L/s). We found that the Upper Snake had surprisingly high biotic demand relative to smaller streams in the same river network for both ammonium (NH4+) and nitrate (NO3-). Placed in the context of a meta-analysis of previously published nutrient uptake studies, these data suggest that large rivers may have similar biotic demand for N as smaller tributaries. We also found that demand for different forms of inorganic N (NH4+ vs. NO3-) scaled differently with stream size. Data from rivers like the Upper Snake and larger are essential for effective water quality management at the scale of river networks. Empirical measurements of solute dynamics in large rivers are needed to understand the role of whole river networks (as opposed to stream reaches) in patterns of nutrient export at regional and continental scales.     

春秋季长江口及其邻近海域营养盐污染研究

依据东海环境监测中心两个航次的资料和美国国家海洋大气管理局(NOAA)相关资料,开发Matlab计算机数值分析和图形显示技术,对长江口及其邻近海域的营养盐污染状况进行分析和探讨。结果表明:海域总体营养盐超标严重;氮、磷污染物的来源主要为径流携带入海,磷还受外海流系高质量浓度磷输入的影响;营养盐污染特征显示,表、底层营养盐均自入海口向外围、由近岸向远岸迅速递减,秋季冲淡水将污染物向外海携带,污染区也相应向外海推移;营养盐结构显示,该区w(N)/w(P)比值最高达到了405,无机氮异常丰富,海域浮游植物生长的限制因子存在时空变化。分析得到营养盐污染特征与长江河口锋位置及冲淡水流向的关系密切,可以考虑由长江河口锋位置及冲淡水流向来确定污染物的聚集位置和扩展方向。     

Unintended nutrient imbalance induced by wastewater effluent inputs to receiving water and its ecological consequences

Eutrophication is the most widespread water quality issue globally. To date, most efforts to control eutrophication have focused on reductions of external nutrient inputs, yet importance of nutrient stoichiometry and subsequent shift in plankton composition in aquatic ecosystem has been largely neglected. To address eutrophication, improved sanitation is one of the United Nations Sustainable Development Goals, spurring the constructions of wastewater treatment facilities that have improved water quality in many lakes and rivers. However, control measures are often targeted at and effective in removing a single nutrient from sewage and thus are less effective in removing the others, resulting in the changes of nutrient stoichiometry. In general, more effective phosphorus removal relative to nitrogen has occurred in wastewater treatment leading to substantial increases in N/P ratios in effluent relative to the influent. Unfortunately, high N/P ratios in receiving waters can impose negative influences on ecosystems. Thus, long-term strategies for domestic wastewater management should not merely focus on the total reduction of nutrient discharge but also consider their stoichiometric balances in receiving waters.     

Marine diatoms grown in chemostats under silicate or ammonium limitation. III. Cellular chemical composition and morphology of Chaetoceros debilis,Skeletonema costatum,and Thalassiosira gravida

Three marine diatoms, Skeletonema costatum, Chaetoceros debilis, and Thalassiosira gravida were grown under no limitation and ammonium or silicate limitation or starvation. Changes in cell morphology were documented with photomicrographs of ammonium and silicate-limited and non-limited cells, and correlated with observed changes in chemical composition. Cultures grown under silicate starvation or limitation showed an increase in particulate carbon, nitrogen and phosporus and chlorophyll a per unit cell volume compared to non-limited cells; particulate silica per cell volume decreased. Si-starved cells were different from Si-limited cells in that the former contained more particulate carbon and silica per cell volume. The most sensitive indicator of silicate limitation or starvation was the ratio C:Si, being 3 to 5 times higher than the values for non-limited cells. The ratios Si:chlorophyll a and S:P were lower and N:Si was higher than non-limited cells by a factor of 2 to 3. The other ratios, C:N, C:P, C:chlorophyll a, N:chlorophyll a, P:chlorophyll a and N:P were considered not to be sensitive indicators of silicate limitation or starvation. Chlorophyll a, and particulate nitrogen per unit cell volume decreased under ammonium limitation and starvation. NH₄-starved cells contained more chlorophyll a, carbon, nitrogen, silica, and phosphorus per cell volume than NH₄-limited cells. N:Si was the most sensitive ratio to ammonium limitation or starvation, being 2 to 3 times lower than non-limited cells. Si:chlorophyll a, P:chlorophyll a and N:P were less sensitive, while the ratios C:N, C:chlorophyll a, N:chlorophyll a, C:Si, C:P and Si:P were the least sensitive. Limited cells had less of the limiting nutrient per unit cell volume than starved cells and more of the non-limiting nutrients (i.e., silica and phosphorus for NH₄-limited cells). This suggests that nutrient-limited cells rather than nutrient-starved cells should be used along with non-limited cells to measure the full range of potential change in cellular chemical composition for one species under nutrient limitation.Contribution No. 943 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Biotic control of stream fluxes: spawning salmon drive nutrient and matter export

Organisms can control movements of nutrients and matter by physically modifying habitat. We examined how an ecosystem engineer, sockeye salmon (Oncorhynchus nerka), influences seasonal fluxes of sediments, nitrogen (N), and phosphorus (P) in streams of southwestern Alaska. The purpose of this study was to investigate whether salmon act as net importers or net exporters of matter and nutrients from streams and how these roles change as a function of salmon population density. We measured discharge and concentrations of suspended sediments and total N and P every 7-14 days for up to four summers in 10 streams spanning a gradient in salmon densities. We statistically allocated whole-season fluxes to salmon activities, such as excretion and bioturbation, and to export by hydrologic discharge. In addition, we used counts of spawning salmon to estimate nutrient and matter imports by salmon to streams. Large seasonal pulses of suspended sediments, P, and N were associated with salmon spawning activities, often increasing export an order of magnitude higher than during pre-salmon levels. Years and streams with more salmon had significantly higher levels of export of sediments and nutrients. In addition, years with higher precipitation had higher background export of P and N. Salmon exported an average of the equivalent of 189%, 60%, and 55% of total matter, P, and N that salmon imported in their bodies. The relative magnitude of export varied; salmon exported more than their bodies imported in 80%, 20%, and 16% across all streams and years for sediments, P, and N, respectively. A bioassay experiment indicated that the P exported by salmon is directly available for use by primary producers in the downstream lake. These results demonstrate that salmon not only move nutrients upstream on large spatial scales via their migration from the ocean and subsequent death, but also redistribute matter and nutrients on finer spatial scales through their spawning activities.     

A model examining hierarchical wetland networks for watershed stormwater management

There is increasing awareness that solutions to degraded quality and excessive quantity of stormwater and resulting impacts on downstream water bodies may require a watershed approach to management rather that the incremental approach that is now common. Examination of low-relief watersheds characteristic of the southeastern coastal plain reveals common hierarchical patterns of surface water convergence that may be emulated in developed watersheds to enhance the efficacy of peak-flow attenuation and pollutant removal. A dynamic systems model was developed to compare stormwater management using a hierarchical network of treatment wetlands with the standard incremental approach wherein treatment systems are designed considering only site-level effluent criteria. The model simulates watershed hydrology, suspended sediment transport and phosphorus removal and transformation. Results indicate that watershed planning of stormwater collection and treatment systems using hierarchical networks can greatly enhance overall effectiveness (annual retention improvements of 31% for flow, 36% for sediment and 27% for phosphorus) with respect to an equal area of uniformly sized wetlands. Further, network proportions can be adjusted to specific runoff characteristics. Distinct roles were observed for each wetland size class: small headwater wetlands effectively removed sediment, medium-sized mid-reach wetlands retained phosphorus, while large wetlands primarily stored and attenuated long-period hydrologic flows.     

Stoichiometry of mesozooplankton in N- and P-limited areas of the Baltic Sea

The Baltic Sea is a very suitable site for stoichiometric studies, since its subbasins differ in their concentration of elemental components, and primary production can therefore be either nitrogen or phosphorus limited. To reveal if the nutrient limitation of mesozooplankton mirrors that of the primary producers, carbon, nitrogen and phosphorus content of both seston and grazers (Acartia sp., Centropages hamatus, Daphnia cristata, Eurytemora affinis, Limnocalanus macrurus, Temora longicornis) were measured in midsummer in the Baltic proper, the Gulf of Finland and the Gulf of Bothnia. The mineral ratios of the different taxa were equal, apart from L. macrurus with notably higher C:P and N:P ratios. Molar C:N ratios were relatively stable (5.1-6.3), whereas C:P and N:P ratios fluctuated more (41-144 and 6.6-24). However, zooplankton elemental composition and limitation did not depend on the limiting nutrient of the phytoplankton, the seston mineral ratio or the sea area. Both the seston-zooplankton elemental imbalance and the food threshold ratio indicated phosphorus limitation of most of the grazers. While L. macrurus may be C or N limited, the possible P deficiency of the other studied taxa suggests that the Baltic Sea zooplankton may act as a potential phosphorus sink, as the freshwater secondary producers do.     

Spatial variation in caddisfly grazing regimes within a northern California watershed

Ecologists seek better understanding of why species interactions change across space and time in natural communities. In streams, species effects on resources and community structure may change as physical characteristics of the stream environment change along drainage networks. We examined spatial and seasonal effects of armored grazers using a small-scale exclusion experiment that was replicated in streams of different drainage areas. Effects of grazing varied with stream size and were related to variation in grazer abundance and phenology. We identified three distinct grazing regimes and a stream size (drainage area [DA]) threshold corresponding to a shift from one to two functional trophic levels. In streams with DA < 1 km2, armored grazers did not reduce biomass of algal biofilms. In slightly larger streams (2-3 km2 DA), the armored grazer guild was dominated by bivoltine Glossosoma. These caddisflies persisted and limited algal biofilms throughout the summer in one of these streams. In the largest tributaries (DA > 10 km2), the grazer guild was dominated by univoltine caddisflies, and grazing limited algal biofilms in early summer, but not late summer, after caddisflies pupated. Drainage area is a useful predictor of spatial transitions in food web interactions within and among watersheds. Quantifying the drainage area threshold at which interactions change in catchments with differing geology, vegetation, hydrology, climate, land use, or species pools should help build the understanding we need to forecast ecological responses to environmental change.