The influence of dissolved nutrients and particulate organic matter quality on microbial respiration and biomass in a forest stream

1. Although dissolved nutrients and the quality of particulate organic matter (POM) influence microbial processes in aquatic systems, these factors have rarely been considered simultaneously. We manipulated dissolved nutrient concentrations and POM type in three contiguous reaches (reference, nitrogen, nitrogen + phosphorus) of a low nutrient, third‐order stream at Hubbard Brook Experimental Forest (U.S.A). In each reach we placed species of leaves (mean C : N of 68 and C : P of 2284) and wood (mean C : N of 721 and C : P of 60 654) that differed in elemental composition. We measured the respiration and biomass of microbes associated with this POM before and after nutrient addition. 2. Before nutrient addition, microbial respiration rates and biomass were higher for leaves than for wood. Respiration rates of microbes associated with wood showed a larger response to increased dissolved nutrient concentrations than respiration rates of microbes associated with leaves, suggesting that the response of microbes to increased dissolved nutrients was influenced by the quality of their substrate. 3. Overall, dissolved nutrients had strong positive effects on microbial respiration and fungal, but not bacterial, biomass, indicating that microbial respiration and fungi were nutrient limited. The concentration of nitrate in the enriched reaches was within the range of natural variation in forest streams, suggesting that natural variation in nitrate among forest streams influences carbon mineralisation and fungal biomass.     

Effects of agriculture on wood breakdown and microbial biofilm respiration in southern Appalachian streams

1. Agriculture causes high sediment, nutrient and light input to streams, which may affect rates of ecosystem processes, such as organic matter decay. In the southern Appalachians, socioeconomic trends over the past 50 years have caused widespread abandonment of farmland with subsequent reforestation. Physical and chemical properties of streams in these reforested areas may be returning to pre‐agriculture levels thereby creating the potential for recovery of ecosystem processes. 2. We examined wood breakdown and microbial activity on wood substrata in streams with different historical and current agricultural activity in their catchments. We analysed historical (1950) and recent (1998) forested land cover from large areas of the southern Appalachians and categorized streams based on percent forested land cover in these two time periods. Categories included a gradient of current agriculture from forested to heavily agricultural and reforestation from agriculture due to land abandonment. We compared microbial respiration on wood veneer substrata and breakdown of wood veneers among these land‐use categories. We also compared temperature, sediment accumulation and nitrogen and phosphorus concentrations. 3. Streams with current agriculture had higher concentrations of dissolved inorganic nitrogen than forested streams. Despite reforestation from agriculture, nitrogen concentrations were also elevated in streams with agricultural histories relative to forested streams. Temperature was also higher in agricultural streams but appeared to recover from historical agriculture through reforestation and stream shading. 4. Wood breakdown rates ranged from 0.0015 to 0.0076 day^?1 and were similar to other studies using wood veneers to determine breakdown rate. Microbial respiration increased with incubation time in streams up to approximately 150 days, after which it remained constant. Neither wood breakdown nor microbial respiration was significantly different among land‐use categories, despite the observed physical and chemical differences in streams based on land‐use. Wood breakdown rates could be predicted by microbial respiration indicating microbial control of wood breakdown in these streams. Both breakdown and microbial respiration were negatively correlated with the amount of inorganic sediment accumulated on wood veneers. 5. Higher nutrients and temperature led us to expect faster breakdown and higher microbial respiration in agricultural streams, but sediment in these streams may be limiting microbial activity and breakdown of organic material resulting in little net effect of agriculture on wood breakdown. Wood may not be desirable as a tool for functional assessment of stream integrity due to its unpredictable response to agriculture.     

Nutrients and temperature additively increase stream microbial respiration

Rising temperatures and nutrient enrichment are co‐occurring global‐change drivers that stimulate microbial respiration of detrital carbon, but nutrient effects on the temperature dependence of respiration in aquatic ecosystems remain uncertain. We measured respiration rates associated with leaf litter, wood, and fine benthic organic matter (FBOM) across seasonal temperature gradients before (PRE) and after (ENR1, ENR2) experimental nutrient (nitrogen [N] and phosphorus [P]) additions to five forest streams. Nitrogen and phosphorus were added at different N:P ratios using increasing concentrations of N (~80–650 μg/L) and corresponding decreasing concentrations of P (~90–11 μg/L). We assessed the temperature dependence, and microbial (i.e., fungal) drivers of detrital mass‐specific respiration rates using the metabolic theory of ecology, before vs. after nutrient enrichment, and across N and P concentrations. Detrital mass‐specific respiration rates increased with temperature, exhibiting comparable activation energies (E, electronvolts [eV]) for all substrates (FBOM E = 0.43 [95% CI = 0.18–0.69] eV, leaf litter E = 0.30 [95% CI = 0.072–0.54] eV, wood E = 0.41 [95% CI = 0.18–0.64] eV) close to predicted MTE values. There was evidence that temperature‐driven increased respiration occurred via increased fungal biomass (wood) or increased fungal biomass‐specific respiration (leaf litter). Respiration rates increased under nutrient‐enriched conditions on leaves (1.32×) and wood (1.38×), but not FBOM. Respiration rates responded weakly to gradients in N or P concentrations, except for positive effects of P on wood respiration. The temperature dependence of respiration was comparable among years and across N or P concentration for all substrates. Responses of leaf litter and wood respiration to temperature and the combined effects of N and P were similar in magnitude. Our data suggest that the temperature dependence of stream microbial respiration is unchanged by nutrient enrichment, and that increased temperature and N + P availability have additive and comparable effects on microbial respiration rates.     

Nutrients stimulate leaf breakdown rates and detritivore biomass: bottom-up effects via heterotrophic pathways

Most nutrient enrichment studies in aquatic systems have focused on autotrophic food webs in systems where primary producers dominate the resource base. We tested the heterotrophic response to long-term nutrient enrichment in a forested, headwater stream. Our study design consisted of 2 years of pretreatment data in a reference and treatment stream and 2 years of continuous nitrogen (N) + phosphorus addition to the treatment stream. Studies were conducted with two leaf species that differed in initial C:N, Rhododendron maximum (rhododendron) and Acer rubrum (red maple). We determined the effects of nutrient addition on detrital resources (leaf breakdown rates, litter C:N and microbial activity) and tested whether nutrient enrichment affected macroinvertebrate consumers via increased biomass. Leaf breakdown rates were ca. 1.5 and 3× faster during the first and second years of enrichment, respectively, in the treatment stream for both leaf types. Microbial respiration rates of both leaf types were 3× higher with enrichment, and macroinvertebrate biomass associated with leaves increased ca. 2–3× with enrichment. The mass of N in macroinvertebrate biomass relative to leaves tended to increase with enrichment up to 6× for red maple and up to 44× for rhododendron leaves. Lower quality (higher C:N) rhododendron leaves exhibited greater changes in leaf nutrient content and macroinvertebrate response to nutrient enrichment than red maple leaves, suggesting a unique response by different leaf species to nutrient enrichment. Nutrient concentrations used in this study were moderate and equivalent to those in streams draining watersheds with altered land use. Thus, our results suggest that similarly moderate levels of enrichment may affect detrital resource quality and subsequently lead to altered energy and nutrient flow in detrital food webs. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Whole-stream nitrate addition affects litter decomposition and associated fungi but not invertebrates

We assessed the effect of whole-stream nitrate enrichment on decomposition of three substrates differing in nutrient quality (alder and oak leaves and balsa veneers) and associated fungi and invertebrates. During the 3-month nitrate enrichment of a headwater stream in central Portugal, litter was incubated in the reference site (mean NO₃-N 82 μg l⁻¹) and four enriched sites along the nitrate gradient (214–983 μg NO₃-N l⁻¹). A similar decomposition experiment was also carried out in the same sites at ambient nutrient conditions the following year (33–104 μg NO₃-N l⁻¹). Decomposition rates and sporulation of aquatic hyphomycetes associated with litter were determined in both experiments, whereas N and P content of litter, associated fungal biomass and invertebrates were followed only during the nitrate addition experiment. Nitrate enrichment stimulated decomposition of oak leaves and balsa veneers, fungal biomass accrual on alder leaves and balsa veneers and sporulation of aquatic hyphomycetes on all substrates. Nitrate concentration in stream water showed a strong asymptotic relationship (Michaelis–Menten-type saturation model) with temperature-adjusted decomposition rates and percentage initial litter mass converted into aquatic hyphomycete conidia for all substrates. Fungal communities did not differ significantly among sites but some species showed substrate preferences. Nevertheless, certain species were sensitive to nitrogen concentration in water by increasing or decreasing their sporulation rate accordingly. N and P content of litter and abundances or richness of litter-associated invertebrates were not affected by nitrate addition. It appears that microbial nitrogen demands can be met at relatively low levels of dissolved nitrate, suggesting that even minor increases in nitrogen in streams due to, e.g., anthropogenic eutrophication may lead to significant shifts in microbial dynamics and ecosystem functioning. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Nitrogen limitation of heterotrophic biofilms in boreal streams

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Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

Comparison of Fungal Activities on Wood and Leaf Litter in Unaltered and Nutrient-Enriched Headwater Streams

Fungi are the dominant organisms decomposing leaf litter in streams and mediating energy transfer to other trophic levels. However, less is known about their role in decomposing submerged wood. This study provides the first estimates of fungal production on wood and compares the importance of fungi in the decomposition of submerged wood versus that of leaves at the ecosystem scale. We determined fungal biomass (ergosterol) and activity associated with randomly collected small wood (<40 mm diameter) and leaves in two southern Appalachian streams (reference and nutrient enriched) over an annual cycle. Fungal production (from rates of radiolabeled acetate incorporation into ergosterol) and microbial respiration on wood (per gram of detrital C) were about an order of magnitude lower than those on leaves. Microbial activity (per gram of C) was significantly higher in the nutrient-enriched stream. Despite a standing crop of wood two to three times higher than that of leaves in both streams, fungal production on an areal basis was lower on wood than on leaves (4.3 and 15.8 g C m⁻² year⁻¹ in the reference stream; 5.5 and 33.1 g C m⁻² year⁻¹ in the enriched stream). However, since the annual input of wood was five times lower than that of leaves, the proportion of organic matter input directly assimilated by fungi was comparable for these substrates (15.4 [wood] and 11.3% [leaves] in the reference stream; 20.0 [wood] and 20.2% [leaves] in the enriched stream). Despite a significantly lower fungal activity on wood than on leaves (per gram of detrital C), fungi can be equally important in processing both leaves and wood in streams.     

Stream acidification increases nitrogen uptake by leaf biofilms: implications at the ecosystem scale

1. While anthropogenic stream acidification is known to lower species diversity and impair decomposition, its effects on nutrient cycling remain unclear. The influence of acid‐stress on microbial physiology can have implications for carbon (C) and nitrogen (N) cycles, linking environmental conditions to ecosystem processes. 2. We collected leaf biofilms from streams spanning a gradient of pH (5.1–6.7), related to chronic acidification, to investigate the relationship between qCO₂ (biomass‐specific respiration; mg CO₂‐C g^?1 fungal C h^?1), a known indicator of stress, and biomass‐specific N uptake (μg NH₄‐N mg^?1 fungal biomass h^?1) at two levels of N availability (25 and 100 μg NH₄‐N L^?1) in experimental microcosms. 3. Strong patterns of increasing qCO₂ (i.e. increasing stress) and increasing microbial N uptake were observed with a decrease in ambient (i.e. chronic) stream pH at both levels of N availability. However, fungal biomass was lower on leaves from more acidic streams, resulting in lower overall respiration and N uptake when rates were standardized by leaf biomass. 4. Results suggest that chronic acidification decreases fungal metabolic efficiency because, under acid conditions, these organisms allocate more resources to maintenance and survival and increase their removal of N, possibly via increased exoenzyme production. At the same time, greater N availability enhanced N uptake without influencing CO₂ production, implying increased growth efficiency. 5. At the ecosystem level, reductions in growth because of chronic acidification reduce microbial biomass and may impair decomposition and N uptake; however, in systems where N is initially scarce, increased N availability may alleviate these effects. Ecosystem response to chronic stressors may be better understood by a greater focus on microbial physiology, coupled elemental cycling, and responses across several scales of investigation.     

Chronic catchment nitrogen enrichment and stoichiometric constraints on the bioavailability of dissolved organic matter from leaf leachate

1. Chronic nitrogen (N) deposition may alter the bioavailability of dissolved organic matter (DOM) in streams by multiple pathways. Elevated N deposition may alter the nutrient stoichiometry of DOM as well as nutrient availability in stream water. 2. We evaluated the influence of a decadal‐scale experimental N enrichment on the relative importance of DOM nutrient content and inorganic nutrient availability on the bioavailability of DOM. We measured the consumption of dissolved organic carbon (DOC) and changes in nutrient concentration, DOM components and enzyme activity in a bottle incubation assay with different DOM and nutrient treatments. To evaluate the effect of DOM stoichiometry, we used leaf leachates of different carbon/N/phosphorus (C : N :P) ratio, made from leaf litter sourced in the reference and N‐enriched catchments at the Bear Brook Watershed in Maine (BBWM). We also manipulated the concentration of inorganic N and P to compare the effect of nutrient enrichment with DOM stoichiometry. 3. DOC from the N‐enriched catchment was consumed 14% faster than that from the reference catchment. However, mean DOC consumption for both leachates was more than doubled by the simultaneous addition of N and P, compared to controls, while the addition of N or P alone increased consumption by 42 and 23%, respectively. The effect of N and/or P enrichment consistently had a greater effect than DOM source for all response variables considered. 4. We subsequently conducted DOC uptake measurements using leaf leachate addition under ambient and elevated N and P in the streams draining the reference and N‐enriched catchments at BBWM. In both streams, DOC uptake lengths were shorter when N and P were elevated. 5. Although both DOM stoichiometry and inorganic nutrient availability affect DOM bioavailability, N and P co‐limitation appears to be the dominant driver of reach‐scale processing of DOM.     

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.     

A meta‐analysis of the effects of nutrient enrichment on litter decomposition in streams

The trophic state of many streams is likely to deteriorate in the future due to the continuing increase in human‐induced nutrient availability. Therefore, it is of fundamental importance to understand how nutrient enrichment affects plant litter decomposition, a key ecosystem‐level process in forest streams. Here, we present a meta‐analysis of 99 studies published between 1970 and 2012 that reported the effects of nutrient enrichment on litter decomposition in running waters. When considering the entire database, which consisted of 840 case studies, nutrient enrichment stimulated litter decomposition rate by approximately 50%. The stimulation was higher when the background nutrient concentrations were low and the magnitude of the nutrient enrichment was high, suggesting that oligotrophic streams are most vulnerable to nutrient enrichment. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher in the laboratory than in the field experiments, suggesting that laboratory experiments overestimate the effect and their results should be interpreted with caution. Among field experiments, effects of nutrient enrichment were smaller in the correlative than in the manipulative experiments since in the former the effects of nutrient enrichment on litter decomposition were likely confounded by other environmental factors, e.g. pollutants other than nutrients commonly found in streams impacted by human activity. However, primary studies addressing the effect of multiple stressors on litter decomposition are still few and thus it was not possible to consider the interaction between factors in this review. In field manipulative experiments, the effect of nutrient enrichment on litter decomposition depended on the scale at which the nutrients were added: stream reach > streamside channel > litter bag. This may have resulted from a more uniform and continuous exposure of microbes and detritivores to nutrient enrichment at the stream‐reach scale. By contrast, nutrient enrichment at the litter‐bag scale, often by using diffusing substrates, does not provide uniform controllable nutrient release at either temporal or spatial scales, suggesting that this approach should be abandoned. In field manipulative experiments, the addition of both nitrogen (N) and phosphorus (P) resulted in stronger stimulation of litter decomposition than the addition of N or P alone, suggesting that there might be nutrient co‐limitation of decomposition in streams. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher for wood than for leaves, and for low‐quality than for high‐quality leaves. The effect of nutrient enrichment on litter decomposition may also depend on climate. The tendency for larger effect size in colder regions suggests that patterns of biogeography of invertebrate decomposers may be modulating the effect of nutrient enrichment on litter decomposition. Although studies in temperate environments were overrepresented in our database, our meta‐analysis suggests that the effect of nutrient enrichment might be strongest in cold oligotrophic streams that depend on low‐quality plant litter inputs.     

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.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Does nitrogen become limiting under high‐P conditions in detritus‐based tropical streams?

1. We examined effects of nutrients on leaf breakdown in interior forest streams at La Selva Biological Station, Costa Rica. We tested the hypothesis that dissolved inorganic nitrogen (DIN) becomes limiting when ambient phosphorus (P) concentration is high. We also compared the breakdown of relatively ‘low quality’ leaves (lower C : N, Trema integerrima) with that of ‘higher quality’ leaves (higher C : N, Ficus insipida) in a high‐P stream. 2. Litterbags were incubated in two streams: one enriched experimentally with P [target concentration 200 μg soluble reactive phosphorus (SRP) L^?1] and one control (naturally low P concentration approximately 10 μg SRP L^?1). Ammonium enrichment was achieved by adding fertiliser upstream of half of the litterbags in each stream. 3. Phosphorus addition stimulated leaf breakdown, microbial respiration, ergosterol and leaf %P. Leaf breakdown rate was consistent with those in La Selva streams with naturally high P concentration. 4. Nitrogen (N) addition had no effect on leaf breakdown, microbial respiration, ergosterol or leaf chemistry in either the P‐enriched or the reference stream, in spite of low N : P ratios. We conclude that N is probably not limiting in streams at La Selva that are naturally high in P. This may be due to moderately high ambient N concentration (>200 μg DIN L^?1) prevailing throughout the year. 5. The species with a lower C : N decomposed more rapidly and supported higher microbial activity than that with a higher C : N. Subtle differences in leaf N content, as well as dissolved P concentration, may be important in determining microbial colonisation and subsequent leaf breakdown.     

The quality of organic matter mediates the response of heterotrophic biofilms to phosphorus enrichment of the water column and substratum

1. Heterotrophic biofilms are important drivers of community respiration, nutrient cycling and decomposition of organic matter in stream ecosystems. Both organic matter quality and nutrient levels have been shown to affect biofilm biomass and activity individually, but both factors have rarely been manipulated simultaneously. 2. To experimentally manipulate the organic matter quality and phosphorus (P) levels of both the substratum and water column, we first used cellulose cloth as a low‐quality organic material and enhanced its quality and P‐content by amending the underlying agar with maltose and P, respectively (Experiment I). To manipulate water column P, artificial substrata were incubated in low‐ and high‐P sites of a whole‐stream P‐enrichment in lowland Costa Rica. 3. Results from Experiment I suggest that heterotrophic biofilm respiration on cellulose cloth is co‐limited by carbon (C) and P. Biofilm respiration responded in an additive manner to combined effects of maltose and P‐enrichment of water column and synergistically to maltose and high‐P in substrata. 4. As decomposing organic matter that supports heterotrophic biofilms varies naturally in its labile C content along with other physical and chemical properties, we conducted a second experiment (Experiment II) in which we amended leaf discs from two species (Trema integerrima, a labile C source and Zygia longifolia, a recalcitrant C source) with maltose. We incubated the substrata in low‐ and high‐P sites of the P‐enrichment stream. 5. Results from Experiment II indicate that biofilm respiration on a labile C source (Trema) was not C‐limited, while biofilm respiration on a recalcitrant C source (Zygia) was C‐limited. Phosphorus stimulated the biofilm respiration and breakdown rate on Trema, but not on Zygia, supporting the hypothesis that the stimulatory effect of P‐enrichment is dependent on the availability of labile C in decomposing leaves. 6. Our results suggest that the interactive effects of organic matter quality and nutrient loading of streams can significantly increase microbial biofilm activity, potentially altering the trophic base of stream food webs. Researchers should consider both the organic matter quality and the enrichment of both water column and substrata to better predict the effects of anthropogenic nutrient loading to stream the ecosystems.     

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

Effects of stream phosphorus levels on microbial respiration

SUMMARY 1. We examined microbial respiration among streams in lowland Costa Rica comprising a natural phosphorus gradient (5–350 μg SRP L^?1) resulting from variable inputs of solute‐rich (e.g. P, SO₄ and Cl) groundwater. 2. Microbial respiration rates were determined by measuring oxygen change in situ in nine low‐order streams on three substrate types: mixed leaves collected from the stream bottom, conditioned Ficus leaves and sediments. 3. Respiration rates on both leaf types were positively related to phosphorus and negatively related to N : P ratios. Microbial respiration rates on sediments were not related to any of the variables [i.e. soluble reactive phosphorus (SRP), N‐NO₃ and N : P] measured. 4. Respiration rates on newly colonised Ficus leaves formed an asymptotic curve increasing to a plateau, suggesting that saturation with phosphorus occurred at concentrations <15 μg SRP L^?1. 5. To test the hypothesis that phosphorus was the main solute in solute‐rich water that was driving observed differences in microbial respiration rates, we artificially enriched a small stream with phosphorus and measured changes in respiration before and after enrichment. 6. Experimental phosphorus enrichment produced increases in respiration rates similar in magnitude to those observed in the nine streams forming the natural phosphorus gradient, supporting our hypothesis that phosphorus was the major variable driving interstream differences in microbial respiration rates. Respiration rates were higher in this study than those reported for most other tropical streams and rivers with the exception of those reported for tropical Asian streams. 7. Results indicate that variations in phosphorus concentrations can potentially affect patterns of microbial respiration rates at a landscape level via differential inputs of solute‐rich groundwater into streams.     

Fungal Activity Associated with Decomposing Wood is Affected by Nitrogen Concentration in Water

We examined the effect of two contrasting nitrogen concentrations in water (0.16 and 0.82 mg L^–1) in the mass loss of pre‐conditioned balsa wood, and associated fungal activity, in a laboratory microcosm experiment. Contrary to our predictions given the poor nutrient quality of balsa wood, an increase in dissolved nitrogen concentration did not result in increased mass loss or microbial oxygen consumption. Conidial production was stimulated in high, but not in low nitrogen microcosms, although the number of fungal species was similar between both treatments. The percentage contribution of each fungal species to total conidial production was similar at both N concentrations. The results support the notion that reproductive activity of aquatic hyphomycetes is the most sensitive microbial parameter to changes in the environment. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Elevated temperature may intensify the positive effects of nutrients on microbial decomposition in streams

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