Experimental nutrient enrichment of forest streams increases energy flow to predators along greener food‐web pathways

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Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.     

Factors affecting ammonium uptake in streams – an inter-biome perspective

1. The Lotic Intersite Nitrogen eXperiment (LINX) was a coordinated study of the relationships between North American biomes and factors governing ammonium uptake in streams. Our objective was to relate inter‐biome variability of ammonium uptake to physical, chemical and biological processes. 2. Data were collected from 11 streams ranging from arctic to tropical and from desert to rainforest. Measurements at each site included physical, hydraulic and chemical characteristics, biological parameters, whole‐stream metabolism and ammonium uptake. Ammonium uptake was measured by injection of ¹⁵N‐ammonium and downstream measurements of ¹⁵N‐ammonium concentration. 3. We found no general, statistically significant relationships that explained the variability in ammonium uptake among sites. However, this approach does not account for the multiple mechanisms of ammonium uptake in streams. When we estimated biological demand for inorganic nitrogen based on our measurements of in‐stream metabolism, we found good correspondence between calculated nitrogen demand and measured assimilative nitrogen uptake. 4. Nitrogen uptake varied little among sites, reflecting metabolic compensation in streams in a variety of distinctly different biomes (autotrophic production is high where allochthonous inputs are relatively low and vice versa). 5. Both autotrophic and heterotrophic metabolism require nitrogen and these biotic processes dominate inorganic nitrogen retention in streams. Factors that affect the relative balance of autotrophic and heterotrophic metabolism indirectly control inorganic nitrogen uptake.     

Effects of upstream lakes and nutrient limitation on periphytic biomass and nitrogen fixation in oligotrophic, subalpine streams

1. We conducted bioassays of nutrient limitation to understand how macronutrients and the position of streams relative to lakes control nitrogen (N₂) fixation and periphytic biomass in three oligotrophic Rocky Mountain catchments. We measured periphytic chlorophyll‐a (chl‐a) and nitrogen‐fixation responses to nitrogen (N) and phosphorus (P) additions using nutrient‐diffusing substrata at 19 stream study sites, located above and below lakes within the study catchments. 2. We found that periphytic chl‐a was significantly co‐limited by N and P at 13 of the 19 sites, with sole limitation by P observed at another four sites, and no nutrient response at the final two sites. On average, the addition of N, P and N + P stimulated chl‐a 35%, 114% and 700% above control values respectively. The addition of P alone stimulated nitrogen fixation by 2500% at five of the 19 sites. The addition of N, either with or without simultaneous P addition, suppressed nitrogen fixation by 73% at nine of the 19 sites. 3. Lake outlet streams were warmer and had higher dissolved organic carbon concentrations than inlet streams and those further upstream, but position relative to lakes did not affect chl‐a and nitrogen fixation in the absence of nutrient additions. Chl‐a response to nutrient additions did not change along the length of the study streams, but nitrogen fixation was suppressed more strongly by N, and stimulated more strongly by P, at lower altitude sites. The responses of chl‐a and nitrogen fixation to nutrients were not affected by location relative to lakes. Some variation in responses to nutrients could be explained by nitrate and/or total N concentration. 4. Periphytic chl‐a and nitrogen fixation were affected by nutrient supply, but responses to nutrients were independent of stream position in the landscape relative to lakes. Understanding interactions between nutrient supply, nitrogen fixation and chl‐a may help predict periphytic responses to future perturbations of oligotrophic streams, such as the deposition of atmospheric N.     

Microbial enzyme activity,nutrient uptake and nutrient limitation in forested streams

1. We measured NH₄⁺ and PO₄^?3 uptake length (S_w), uptake velocity (V_f), uptake rate (U), biofilm respiration and enzyme activity and channel geomorphology in streams draining forested catchments in the northwestern (Northern California Coast Range and Cascade Mountains) and southeastern (Appalachian and Ouachita mountains) regions of the United States. Our goal was to use measures of biofilm enzyme activity and nutrient uptake to assess nutrient limitation in forested streams across broad regional scales. 2. Geomorphological attributes, biofilm enzyme activity and NH₄⁺ uptake were significantly different among streams in the four study units. There was no study unit effect on PO₄^?3 uptake. The proportion of the stream channel in pools, % woody debris, % canopy closure, median substrate size (d₅₀), stream width (w), stream velocity (v), discharge (Q), dispersion coefficient (D) and transient storage (A_s/A) were correlated with biofilm enzyme activity and nutrient uptake in some study units. 3. Canonical correlation analyses across study units revealed significant correlations of NH₄‐V_f and PO₄‐V_f with geomorphological attributes (w, d₅₀, D, % woody debris, channel slope and % pools) and biofilm phosphatase activity. 4. The results did not support our expectation that carbon processing rates by biofilm microbial assemblages would be governed by stream nutrient availability or that resulting biofilm enzyme activity would be an indicator of nutrient uptake. However, the relative abundances of peptidases, phosphatase and glycosidases did yield insight into potential N‐, P‐ and C‐limitation of stream biofilm assemblages, and our use of biofilm enzyme activity represents a novel application for understanding nutrient limitations in forested streams. 5. Regressions of V_f and U against ambient NH₄⁺ and PO₄^?3 indicated that none of our study streams was either NH₄⁺ or PO₄^?3 saturated. The Appalachian, Ouachita and Coastal streams showed evidence of NH₄⁺ limitation; the Ouachita and Coastal streams were PO₄^?3 limited. As a correlate of nutrient limitation and saturation in streams, ratios of total aminopeptidase and phosphatase activities and the ratio of NH₄‐U to PO₄‐U indicate these forested streams are predominantly N‐limited, with only the streams draining Ouachita and Coastal catchments demonstrating appreciable levels of P‐limitation. 6. Our results comparing the stoichiometry of microbial enzyme activity with nutrient uptake ratios and with the molar ratios N and P in stream waters suggest that biological limitations are not strictly the result of stream chemistry and that the assessments of nutrient limitations in stream ecosystems should not be based on chemistry alone. 7. Our present study, along with previous work in streams, rivers and wetlands, suggests that microbial enzyme activities, especially the ratios of total peptidases to phosphatase, are useful indicators of nutrient limitations in aquatic ecosystems.     

Reproductive patterns and secondary production of <Emphasis Type="Italic">Gammaropsis japonicus</Emphasis> (Crustacea,Amphipoda) on the seagrass <Emphasis Type="Italic">Zostera marina</Emphasis> of Korea

It is essential to know the nutrient limitation status of biofilms to understand how they may buffer uptake and export of nutrients from polluted watersheds. We tested the effects of nutrient additions on biofilm biomass (chlorophyll a, ash free dry mass (AFDM), and autotrophic index (AI, AFDM/chl a)) and metabolism via nutrient-diffusing substrate bioassays (control, nitrogen (N), phosphorus (P), and N + P treatments) at 11 sites in the Upper Snake River basin (southeast Idaho, USA) that differed in the magnitude and extent of human-caused impacts. Water temperature, turbidity, and dissolved inorganic N concentrations all changed seasonally at the study sites, while turbidity and dissolved inorganic N and P also varied with impact level. Chl a and AI on control treatments suggested that the most heavily impacted sites supported more autotrophic biofilms than less-impacted sites, and that across all sites biofilms were more heterotrophic in autumn than in summer. Nutrient stimulation or suppression of biofilm biomass was observed for chl a in 59% of the experiments and for AFDM in 33%, and the most frequent response noted across all study sites was N limitation. P suppression of chl a was observed only at the most-impacted sites, while AFDM was never suppressed by nutrients. When nutrient additions did have significant effects on metabolism, they were driven by differences in biomass rather than by changes in metabolic rates. Our study demonstrated that biofilms in southeast Idaho rivers were primarily limited by N, but nutrient limitation was more frequent at sites with good water quality than at those with poor water quality. Additionally, heterotrophic and autotrophic biofilm components may respond differently to nutrient enrichment, and nutrient limitation of biofilm biomass should not be considered a surrogate for metabolism in these rivers. Handling editor: D. Ryder     

Increased resource use efficiency amplifies positive response of aquatic primary production to experimental warming

Climate warming is affecting the structure and function of river ecosystems, including their role in transforming and transporting carbon (C), nitrogen (N), and phosphorus (P). Predicting how river ecosystems respond to warming has been hindered by a dearth of information about how otherwise well‐studied physiological responses to temperature scale from organismal to ecosystem levels. We conducted an ecosystem‐level temperature manipulation to quantify how coupling of stream ecosystem metabolism and nutrient uptake responded to a realistic warming scenario. A ~3.3°C increase in mean water temperature altered coupling of C, N, and P fluxes in ways inconsistent with single‐species laboratory experiments. Net primary production tripled during the year of experimental warming, while whole‐stream N and P uptake rates did not change, resulting in 289% and 281% increases in autotrophic dissolved inorganic N and P use efficiency (UE), respectively. Increased ecosystem production was a product of unexpectedly large increases in mass‐specific net primary production and autotroph biomass, supported by (i) combined increases in resource availability (via N mineralization and N₂ fixation) and (ii) elevated resource use efficiency, the latter associated with changes in community structure. These large changes in C and nutrient cycling could not have been predicted from the physiological effects of temperature alone. Our experiment provides clear ecosystem‐level evidence that warming can shift the balance between C and nutrient cycling in rivers, demonstrating that warming will alter the important role of in‐stream processes in C, N, and P transformations. Moreover, our results reveal a key role for nutrient supply and use efficiency in mediating responses of primary producers to climate warming.     

Functional responses of stream biofilms to flow cessation,desiccation and rewetting

1. Mediterranean climate regions are characterised by long summer droughts that usually involve flow intermittency in low‐ to mid‐order streams. Flow intermittency implies flow cessation, drying and subsequent rewetting of the streambed, and affects both autotrophic and heterotrophic processes. The balance between these processes, as well as the balance in the use of carbon (C), nitrogen (N) and phosphorus (P) may change because of the ongoing increase in stream flow intermittency caused by global change in many regions. It is therefore crucial to understand better the consequences of this phenomenon. 2. Our two initial hypotheses were (i) that flow intermittency would impact more on autotrophic than on heterotrophic processes in stream biofilms owing to the higher water dependence of autotrophs, as well as differences in the water storage capacity of the stream biofilm compartments where autotrophic and heterotrophic processes mainly occur (surface cobbles versus hyporheic sediments) and (ii) that the C‐N‐P use by biofilms would change during the dry period (terrestrial phase) owing to the extreme water stress conditions. These hypotheses were tested by analysing the functional response of the main stream biofilms (epilithic, epipsammic and hyporheic) during flow cessation, desiccation and rewetting in a Mediterranean forested stream. The autotrophic response was characterised through changes in the photon yield, whereas the heterotrophic response was characterised by changes in the extracellular enzyme activities. 3. Streambed desiccation had clear effects on the functioning of stream biofilms. Autotrophic biomass decreased by 80% with streambed desiccation, but recovered rapidly after flow resumption. Heterotrophs were more resistant to water stress, especially in the epipsammic and hyporheic biofilms where bacterial cell density decreased only by 20%. 4. Extracellular enzyme activities remained relatively high, and the balance in the C‐N‐P use by biofilms changed during the dry period. The C and P breakdown capacities were maintained during dry conditions, especially in the epipsammic and hyporheic biofilms, but the degradation of N compounds sharply decreased. Elemental molar ratios (C:N and C:P) of the different biofilms also changed with streambed desiccation. C:P ratios increased from 80 to 300, while the C:N ratios increased from 10 to 16. 5. Given the contrasting responses of autotrophic and heterotrophic processes in the different biofilms, our results suggest that the current increase in flow intermittency extent is likely to increase the relative importance of heterotrophic processes in stream ecosystems, as well as the relative contribution of the hyporheic biofilm to C‐N‐P use. Our results further suggest that the longer streams remain dry, the more the biofilm stoichiometry will change.     

Nutrient limitation of epilithic and epixylic biofilms in ten North American streams

1. Nutrient diffusing substrata were used to determine the effect of added inorganic nitrogen (N) and phosphorus (P) on the development of epilithic and epixylic biofilms in 10 North American streams. Four treatments of diffusing substrata were used: Control (agar only), N addition (0.5 m NaNO₃), P addition (0.5 m KH₂PO₄), and N + P combined (0.5 m NaNO₃ + 0.5 m KH₂PO₄). Agar surfaces were covered with glass fibre filters (for epilithon) or discs of untreated white oak wood veneer (for epixylon). 2. We found that if algae showed significant response to nutrient addition, N limitation (either N alone or N with P) was the most frequent response both on GF/F filters and on wood. Despite the low dissolved nutrient concentrations in our study streams, more than a third of the streams did not show any response to N or P addition. In fact, P was never the sole limiting nutrient for algal biofilms in this study. 3. Nutrient addition influenced algal colonisation of inorganic versus organic substrata in different ways. The presence of other biofilm constituents (e.g. fungi or bacteria) may influence whether algal biomass on wood increased in response to nutrient addition. Algae on organic and inorganic substrata responded similarly to nutrient addition in only one stream. 4. Fungal biomass on wood was nutrient limited in six of 10 study streams. N limitation of fungal biomass (with or without secondary P limitation) was most frequent, but P limitation did occur in two streams. 5. Our results show that biomass responses to nutrient addition by the heterotrophic and autotrophic components of the epixylic biofilm were different, though both experienced the same stream nutrient conditions. For algae and fungi growing on wood, limiting nutrients were rarely similar. Only three of nine streams showed the same biomass response to nutrient addition, including two that showed no significant change in biomass despite added nutrients.     

Persistent nitrogen limitation of stream biofilm communities along climate gradients in the Arctic

Climate change is rapidly reshaping Arctic landscapes through shifts in vegetation cover and productivity, soil resource mobilization, and hydrological regimes. The implications of these changes for stream ecosystems and food webs is unclear and will depend largely on microbial biofilm responses to concurrent shifts in temperature, light, and resource supply from land. To study those responses, we used nutrient diffusing substrates to manipulate resource supply to biofilm communities along regional gradients in stream temperature, riparian shading, and dissolved organic carbon (DOC) loading in Arctic Sweden. We found strong nitrogen (N) limitation across this gradient for gross primary production, community respiration and chlorophyll‐a accumulation. For unamended biofilms, activity and biomass accrual were not closely related to any single physical or chemical driver across this region. However, the magnitude of biofilm response to N addition was: in tundra streams, biofilm response was constrained by thermal regimes, whereas variation in light availability regulated this response in birch and coniferous forest streams. Furthermore, heterotrophic responses to experimental N addition increased across the region with greater stream water concentrations of DOC relative to inorganic N. Thus, future shifts in resource supply to these ecosystems are likely to interact with other concurrent environmental changes to regulate stream productivity. Indeed, our results suggest that in the absence of increased nutrient inputs, Arctic streams will be less sensitive to future changes in other habitat variables such as temperature and DOC loading.     

Effects of irradiance on diel and seasonal patterns of nutrient uptake by stream periphyton

1. Irradiance strongly affects the abundance of stream periphyton communities that in turn influence patterns of instream nutrient uptake. We examined relationships between irradiance and periphyton nutrient uptake taking into account diel and seasonal variation in ambient irradiance. 2. Uptake of dissolved N, P and C by periphyton as areal uptake (U) and demand (V_f) was determined under 11 irradiance levels (0–100% of ambient conditions) using shallow stream‐side experimental channels. Experiments were conducted once per season over one annual cycle with both day and night uptake rates assessed, together with periphyton biomass and autotrophic production rates. 3. No consistent diel variation in areal uptake or demand was detected for the predominant inorganic or total dissolved nutrients even at the highest irradiances. Lack of variation may indicate nutrient limitation, with photosynthetic sequestration and storage of C during the day for subsequent utilisation at night. Alternatively, oxygen consumption by photoautotrophs at night may stimulate compensatory heterotrophic uptake (e.g. denitrification). 4. In all seasons, release of dissolved organic N was detected during the day but to a lesser extent at night. This was not directly related to irradiance levels, indicating that heterotrophic metabolism (e.g. microbial decomposition) contributes to this phenomenon. 5. Areal uptake and demand for the predominant inorganic and total dissolved nutrients increased in response to increasing irradiance in some or all seasons, but rates were typically higher during the spring and summer. Saturation of areal uptake and demand at elevated irradiances was evident during the spring. demand was also saturated at higher irradiances in the summer and autumn. Maximum demand was comparable during spring and summer, but saturation occurred at lower irradiance in summer (24 h average 135–145 μmol m^?2 s^?1) relative to spring (312–424 μmol m^?2 s^?1), indicating more efficient nutrient uptake in summer. Higher total periphyton biomass in summer, but comparable autotrophic biomass (chlorophyll a), implies that heterotrophic metabolism may contribute to this greater efficiency. In spring, autotrophic biomass peaked at an irradiance level of 225 μmol m^?2 s^?1, also suggesting a role for heterotrophic metabolism in demand at higher irradiances. 6. The results of this study show that irradiance levels exert a strong influence on the nature and quantity of instream nutrient uptake with N demand saturated at elevated irradiance levels during the spring, summer and autumn. Our results also suggest that heterotrophic metabolism makes a measurable contribution to instream nutrient uptake even under higher irradiances that favour autotrophic activity.     

Regulation of phosphate uptake reveals cyanobacterial bloom resilience to shifting N:P ratios

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Effects of nutrients and light on periphyton biomass and nitrogen uptake in Mediterranean streams with contrasting land uses

1. Nutrient diffusing substrata (NDS) were used to determine the relative importance of nutrients and light as potential limiting factors of periphyton biomass and nitrogen (N) uptake in Mediterranean streams subjected to different human impacts. The nutrients examined were phosphorus (P) and N, and we also further differentiated between the response of periphyton communities to N species (i.e. NO₃‐N and NH₄‐N). To examine the effect of light and nutrients on periphyton biomass, chlorophyll a accrual rates on NDS located at open and closed canopy sites were compared. The effect of nutrient availability on periphyton uptake was measured by ¹⁵N changes on the NDS after NO₃‐¹⁵N short‐term nutrient additions. 2. Results show that light was the main factor affecting algal biomass in the study streams. Algal biomass was in general higher at open than at closed canopy sites. Nutrient availability, as simulated with the NDS experiments, did not enhance algal biomass accrual in either of the 2 light conditions. 3. In the control treatments (i.e. ambient concentrations), periphyton NO₃‐N uptake rates increased and C : N molar ratios decreased consistently with increases in N availability across streams. NO₃‐N uptake rates were altered when ambient N concentrations were increased artificially in the N amended NDS. Periphyton assemblages growing on N enriched substrata seemed to preferentially take up N diffusing from the substratum rather than N from the water column. This response differed among streams, and depended on ambient N availability. 4. Periphyton biomass was not significantly different between substrata exposed to the two forms of available N sources. Nonetheless, we found differences in the effects of both N sources on the uptake of N from the water column. NH₄‐N seemed to be the preferred source of N for periphyton growing on NDS. 5. Results suggest that the effect of riparian zones on light availability, although seldom considered by water managers, may be more important than nutrients in controlling eutrophication effects derived from human activities. Finally, our results confirm that not only increases in concentration, but also stoichiometric imbalances should be considered when examining N retention in human altered streams.     

Leaf litter identity alters the timing of lotic nutrient dynamics

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Phosphorus Regeneration in Fresh-Water Paramecia*

Turnover rates of soluble phosphorus (P) by Paramecium multimicronucleatum were compared under 3 temperature regimes. Phosphorus uptake by this heterotroph was almost completely dependent on consumption of bacteria rather than direct uptake of soluble ³²PO₄. Phosphorus uptake rates by P. bursaria when in its autotrophic state (a symbiotic relationship with zoochlorellae) and when in its heterotrophic (bacteria feeding) state were also compared. Paramecium bursaria showed a much higher uptake of P when large numbers of labeled bacteria were present than when they were absent. Phosphorus turnover rates by P. bursaria were determined and compared with those of P. multimicronucleatum. These ciliates showed remarkably rapid turnover rates for soluble P. In each case turnover rates were universely related to biomass. Ciliates, although small, have very high metabolic rates and therefore may be important for nutrient regeneration in our fresh-water lakes, streams, and ponds.     

Imbalanced nutrient recycling in a warmer ocean driven by differential response of extracellular enzymatic activities

Ocean oligotrophication concurrent with warming weakens the capacity of marine primary producers to support marine food webs and act as a CO₂ sink, and is believed to result from reduced nutrient inputs associated to the stabilization of the thermocline. However, nutrient supply in the oligotrophic ocean is largely dependent on the recycling of organic matter. This involves hydrolytic processes catalyzed by extracellular enzymes released by bacteria, which temperature dependence has not yet been evaluated. Here, we report a global assessment of the temperature‐sensitivity, as represented by the activation energies (E_a), of extracellular β‐glucosidase (βG), leucine aminopeptidase (LAP) and alkaline phosphatase (AP) enzymatic activities, which enable the uptake by bacteria of substrates rich in carbon, nitrogen, and phosphorus, respectively. These E_a were calculated from two different approaches, temperature experimental manipulations and a space‐for‐time substitution approach, which generated congruent results. The three activities showed contrasting E_a in the subtropical and tropical ocean, with βG increasing the fastest with warming, followed by LAP, while AP showed the smallest increase. The estimated activation energies predict that the hydrolysis products under projected warming scenarios will have higher C:N, C:P and N:P molar ratios than those currently generated, and suggest that the warming of oceanic surface waters leads to a decline in the nutrient supply to the microbial heterotrophic community relative to that of carbon, particularly so for phosphorus, slowing down nutrient recycling and contributing to further ocean oligotrophication.     

Increased N affects P uptake of eight grassland species: the role of root surface phosphatase activity

Increased N deposition may change species composition in grassland communities by shifting them to P limitation. Interspecific differences in P uptake traits might be a crucial yet poorly understood factor in determining the N effects. To test the effects of increased N supply (relative to P), we conducted two greenhouse fertilization experiments with eight species from two functional groups (grasses, herbs), including those common in P and N limited grasslands. We investigated plant growth and P uptake from two P sources, orthophosphate and not‐readily available P (bound‐P), under different N supply levels. Furthermore, to test if the N effects on P uptake was due to N availability alone or altered N:P ratio, we examined several uptake traits (root‐surface phosphatase activity, specific root length (SRL), root mass ratio (RMR)) under varying N:P supply ratios. Only a few species (M. caerulea, A. capillaris, S. pratensis) could take up a similar amount of P from bound‐P to that from orthophosphate. These species had neither higher SRL, RMR, phosphatase activity per unit root (Pase_root), nor higher total phosphatase activity (Pase_tot: Pase_root times root mass), but higher relative phosphatase activity (Pase_rel: Pase_tot divided by biomass) than other species. The species common from P‐limited grasslands had high Pase_rel. P uptake from bound‐P was positively correlated with Pase_tot for grasses. High N supply stimulated phosphatase activity but decreased RMR and SRL, resulting in no increase in P uptake from bound‐P. Pase_root was influenced by N:P supply ratio, rather than by only N or P level, whereas SRL and RMR was not dominantly influenced by N:P ratio. We conclude that increased N stimulates phosphatase activity via N:P stoichiometry effects, which potentially increases plant P uptake in a species‐specific way. N deposition, therefore, may alter plant community structure not only by enhancing productivity, but also by favouring species with traits that enable them to persist better under P limited conditions.     

Grazer species effects on epilithon nutrient composition

1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no‐grazer treatment. 4. Traditionally, studies have focused on how grazer‐induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.     

Effects of consumers and nitrogen availability on heterotrophic microbial activity during leaf decomposition in headwater streams

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Bromeliad growth and stoichiometry: responses to atmospheric nutrient supply in fog-dependent ecosystems of the hyper-arid Atacama Desert, Chile

Carbon, nitrogen, and phosphorus (C, N, P) stoichiometry influences the growth of plants and nutrient cycling within ecosystems. Indeed, elemental ratios are used as an index for functional differences between plants and their responses to natural or anthropogenic variations in nutrient supply. We investigated the variation in growth and elemental content of the rootless terrestrial bromeliad Tillandsia landbeckii, which obtains its moisture, and likely its nutrients, from coastal fogs in the Atacama Desert. We assessed (1) how fog nutrient supply influences plant growth and stoichiometry and (2) the response of plant growth and stoichiometry to variations in nutrient supply by using reciprocal transplants. We hypothesized that T. landbeckii should exhibit physiological and biochemical plastic responses commensurate with nutrient supply from atmospheric deposition. In the case of the Atacama Desert, nutrient supply from fog is variable over space and time, which suggests a relatively high variation in the growth and elemental content of atmospheric bromeliads. We found that the nutrient content of T. landbeckii showed high spatio-temporal variability, driven partially by fog nutrient deposition but also by plant growth rates. Reciprocal transplant experiments showed that transplanted individuals converged to similar nutrient content, growth rates, and leaf production of resident plants at each site, reflecting local nutrient availability. Although plant nutrient content did not exactly match the relative supply of N and P, our results suggest that atmospheric nutrient supply is a dominant driver of plant growth and stoichiometry. In fact, our results indicate that N uptake by T. landbeckii plants depends more on N supplied by fog, whereas P uptake is mainly regulated by within-plant nutrient demand for growth. Overall, these findings indicate that variation in fog nutrient supply exerts a strong control over growth and nutrient dynamics of atmospheric plants, which are ubiquitous across fog-dominated ecosystems.