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.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

Do leaf-litter decomposers control biofilm primary production and benthic algal community structure in forest streams? Insights from an outdoor mesocosm experiment

• 1. Forested headwater streams are generally considered to be light-limited ecosystems where primary production is reduced, and the main source of energy and nutrients is composed of allochthonous detritus. We hypothesised that in these ecosystems, the development of primary producers might also be limited by (1) competition for nutrients with leaf-litter decomposers (e.g. bacteria and fungi), and (2) leaf-litter leachates or allelopathic compounds produced by aquatic fungi.
• 2. To test these hypotheses, a 48-day mesocosm experiment was performed in 12 artificial streams containing stream water inoculated with epilithic biofilm suspensions collected from a forested headwater stream. Three different treatments were applied: control without leaf litter (C), microbially conditioned leaf litter added at the beginning of the experiment and left to decompose throughout the experiment (L), or leaf litter renewed three times during the experiment (RL).
• 3. We predicted that (1) the presence of litter, through microbial nutrient immobilisation and allelopathy, would reduce primary production and that (2) this effect would be amplified by litter renewal. We also predicted that nutrient competition would mean that (3) leaf-litter decomposers will alter primary producer community composition and physiology. These predictions were tested by analysing biofilm development, physiology, stoichiometry, and benthic algal community structure. To distinguish between the effects of nutrient immobilisation and allelopathy, the biofilm responses to leaf-litter leachates collected after different microbial conditioning durations were also measured in a parallel laboratory experiment.
• 4. Contrary to our expectations, by day 28, primary producer growth was higher in the mesocosms containing leaf litter (L and RL) despite the rapid decrease in dissolved nutrients when leaf litter was present. After 48 days, the lowest phototrophic biofilm development was observed when leaf litter was renewed (RL), whereas phototrophic biofilm development was similar in the C and L treatments. Biofilm stoichiometry indicated that this effect was most probably related to greater nitrogen limitation in the RL treatment. The presence of leaf litter also affected primary producers' photophysiology, which could be attributed to changes in taxonomic composition and to physiological adjustments of primary producers.
• 5. Laboratory measurements showed that despite a strong inhibition of primary producer growth by unconditioned leaf-litter leachates, microbially conditioned leaf litter had either low or no effects on the development of primary producers.
• 6. These results reveal that leaf-litter decomposers can have both positive and negative effects on primary producers underlining the need to consider microbial interactions when investigating the functioning of forested headwater streams.

     

Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

Effects of Fungal Inocula and Habitat Conditions on Alder and Eucalyptus Leaf Litter Decomposition in Streams of Northern Spain

We investigated how fungal decomposer (aquatic hyphomycetes) communities colonizing alder and eucalyptus leaf litter respond to changes in habitat characteristics (transplantation experiment). We examined the breakdown of leaf materials and the associated fungal communities at two contrasting sites, a headwater stream (H) and a midreach (M). Agroforestry increased from headwater to midreach. One month after the start of experiments at both sites, some leaf samples from the midreach site were transplanted to the headwater site (M–H treatment). Although both sites showed similar dissolved inorganic nutrient concentrations, eucalyptus leaves initially incubated at the midreach site (M, M–H) increased their breakdown rate compared to those incubated along the experiment at the headwater site (H). Alder breakdown rate was not enhanced, suggesting that their consumption was not limited by nutrient availability. Sporulation rates clearly differed between leaf types (alder > eucalyptus) and streams (H > M), but no transplantation effect was detected. When comparing conidial assemblages after transplantation, an inoculum effect (persistence of early colonizing species) was clear in both leaf species. Substrate preference and shifts in the relative importance of some fungal species along the process were also observed. Overall, our results support the determining role of the initial conditioning phase on the whole litter breakdown process, highlighting the importance of intrinsic leaf characteristics and those of the incubation habitat.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

Contrasting nutrient exports from a forested and an agricultural catchment in south-eastern Australia

Dissolved organic carbon (DOC) and total and inorganic nitrogen and phosphorus concentrations were determined over 3 years in headwater streams draining two adjacent catchments. The catchments are currently under different land use; pasture/grazing vs plantation forestry. The objectives of the work were to quantify C and nutrient export from these landuses and elucidate the factors regulating export. In both catchments, stream water dissolved inorganic nutrient concentrations exhibited strong seasonal variations. Concentrations were highest during runoff events in late summer and autumn and rapidly declined as discharge increased during winter and spring. The annual variation of stream water N and P concentrations indicated that these nutrients accumulated in the catchments during dry summer periods and were flushed to the streams during autumn storm events. By contrast, stream water DOC concentrations did not exhibit seasonal variation. Higher DOC and NO₃ ⁻ concentrations were observed in the stream of the forest catchment, reflecting greater input and subsequent breakdown of leaf-litter in the forest catchment. Annual export of DOC was lower from the forested catchment due to the reduced discharge from this catchment. In contrast however, annual export of nitrate was higher from the forest catchment suggesting that there was an additional NO₃ ⁻ source or reduction of a NO₃ ⁻ sink. We hypothesize that the denitrification capacity of the forested catchment has been significantly reduced as a consequence of increased evapotranspiration and subsequent decrease in streamflow and associated reduction in the near stream saturated area.     

Carbon, nitrogen, and phosphorus dynamics during leaf decay in nutrient-enriched stream microecosystems

We investigated the effects of long-term enrichment with nitrate, phosphate, and nitrate+phosphate on the first 5 weeks of leaf detritus processing in laboratory stream microecosystems. Enrichment with nitrate+phosphate accelerated leaf weight loss and increased rates of respiration associated with the leaves. However, whole-system respiration was little changed from that observed in the control stream since respiration in the water was greatly reduced. Enrichment with phosphate alone had little effect except to lower respiration associated with leaf discs. Enrichment with nitrate alone also decreased leaf-disc respiration but resulted in a greatly increased rate of respiration in the water. Net leaching and fragmentation of carbon from the leaves was also increased by nitrate enrichment. Nitrogen and phosphorus levels in leaf material were little affected by enrichment with nitrate or phosphorus alone. Leaves in those streams and in the control stream released nitrogen and phosphorus to the water. In contrast, percent nitrogen and phosphorus increased greatly in the leaves in the stream enriched with both nitrate and phosphate. The leaves in this system immobilized both nitrogen and phosphorus from the water. We also studied the importance of nitrogen fixation as a vector for nitrogen incorporation associated with leaf decomposition in streams. Somewhat surprisingly, fixation by microbes associated with the leaves and by microbes suspended in the water occurred under all three experimental enrichment treatments as well as in the control, casting doubt on the effectiveness of nitrate in inhibiting nitrogenase synthesis in nature. However, N₂-fixation is only a minor source of nitrogen for leaves decaying under the conditions studied.     

Effects of nutrient enrichment on the decomposition of wood and associated microbial activity in streams

1. We determined the effects of nutrient enrichment on wood decomposition rates and microbial activity during a 3‐year study in two headwater streams at Coweeta Hydrologic Laboratory, NC, U.S.A. After a 1‐year pretreatment period, one of the streams was continuously enriched with inorganic nutrients (nitrogen and phosphorus) for 2 years while the other stream served as a reference. We determined the effects of enrichment on both wood veneers and sticks, which have similar carbon quality but differ in physical characteristics (e.g. surface area to volume ratios, presence of bark) that potentially affect microbial colonisation and activity. 2. Oak wood veneers (0.5 mm thick) were placed in streams monthly and allowed to decompose for approximately 90 days. Nutrient addition stimulated ash‐free dry mass loss and increased mean nitrogen content, fungal biomass and microbial respiration on veneers in the treatment stream compared with the reference. The magnitude of the response to enrichment was great, with mass loss 6.1 times, and per cent N, fungal biomass and microbial respiration approximately four times greater in the treatment versus reference stream. 3. Decomposition rate and nitrogen content of maple sticks (ca. 1–2 cm diameter) also increased; however, the effect was less pronounced than for veneers. Wood response overall was greater than that determined for leaves in a comparable study, supporting the hypothesis that response to enrichment may be greater for lower quality organic matter (high C : N) than for higher quality (low C : N) substrates. 4. Our results show that moderate nutrient enrichment can profoundly affect decomposition rate and microbial activity on wood in streams. Thus, the timing and availability of wood that provides retention, structure, attachment sites and food in stream ecosystems may be affected by nutrient concentrations raised by human activities.     

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.     

Water abstraction impacts stream ecosystem functioning via wetted‐channel contraction

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Macroinvertebrate diversity in headwater streams: a review

1. Headwater streams are ubiquitous in the landscape and are important sources of water, sediments and biota for downstream reaches. They are critical sites for organic matter processing and nutrient cycling, and may be vital for maintaining the 'health' of whole river networks.
2. Macroinvertebrates are an important component of biodiversity in stream ecosystems and studies of macroinvertebrate diversity in headwater streams have mostly viewed stream systems as linear reaches rather than as networks, although the latter may be more appropriate to the study of diversity patterns in headwater systems.
3. Studies of macroinvertebrate diversity in headwater streams from around the world illustrated that taxonomic richness is highly variable among continents and regions, and studies addressing longitudinal changes in taxonomic richness of macroinvertebrates generally found highest richness in mid-order streams.
4. When stream systems are viewed as networks at the landscape-scale, α-diversity may be low in individual headwater streams but high β-diversity among headwater streams within catchments and among catchments may generate high γ-diversity.
5. Differing ability and opportunity for dispersal of macroinvertebrates, great physical habitat heterogeneity in headwater streams, and a wide range in local environmental conditions may all contribute to high β-diversity among headwater streams both within and among catchments.
6. Moving beyond linear conceptual models of stream ecosystems to consider the role that spatial structure of river networks might play in determining diversity patterns at the landscape scale is a promising avenue for future research.     

Leaf Litter as a Source of Dissolved Organic Carbon in Streams

Dissolved organic carbon (DOC) is an abundant form of organic matter in stream ecosystems. Most research has focused on the watershed as the source of DOC in streams, but DOC also comes from leaching of organic matter stored in the stream channel. We used a whole-ecosystem experimental approach to assess the significance of leaching of organic matter in the channel as a source of DOC in a headwater stream. Inputs of leaf litter were excluded from a forested Appalachian headwater stream for 3 years. Stream-water concentration, export, and instream generation of DOC were reduced in the litter-excluded stream as compared with a nearby untreated reference stream. The proportion of high molecular weight (HMW) DOC (more than 10,000 daltons) in stream water was not altered by litter exclusion. Mean DOC concentration in stream water was directly related to benthic leaf-litter standing stock. Instream generation of DOC from leaf litter stored in the stream channel contributes approximately 30% of daily DOC exports in this forested headwater stream. This source of DOC is greatest during autumn and winter and least during spring and summer. It is higher during increasing discharge than during base flow. We conclude that elimination of litter inputs from a forested headwater stream has altered the biogeochemistry of DOC in this ecosystem. Received 2 September 1997; accepted 27 January 1998.     

Controls on periphyton biomass in heterotrophic streams

1. Headwater streams of the Hubbard Brook Experimental Forest (HBEF) are typically characterised by a periphyton assemblage of low biomass and diversity. However, periphyton blooms have been observed following catchment deforestation experiments and occasionally during the annual spring thaw before canopy leaf‐out. 2. There is pronounced seasonal variation in both nutrient and light availability in HBEF streams. Stream water nitrogen (N) concentrations and light levels are higher before canopy leaf‐out and after leaf senescence and are lower during the growing season. Periphyton accrual rates also change seasonally; they are highest in spring prior to leaf‐out and significantly lower during summer and in autumn. 3. Periphyton biomass rarely responded positively to in‐situ experimental enrichment with nitrogen or phosphorus. In the summer, nutrient enrichment overall had no effect on periphyton biomass, while outside the growing season N enrichment had inhibitory effects on periphyton. 4. Despite these experimental results, surveys of ambient chlorophyll a concentrations in streams across the HBEF demonstrated no relationship between streamwater dissolved inorganic N or P concentrations and benthic chlorophyll a. 5. Our results suggest that HBEF periphyton communities are not closely regulated by nutrient availability, even during periods of high light availability. The inhibitory effects of nutrient enrichment outside the growing season are interesting, but further research is necessary to elucidate the mechanisms driving these responses.     

The Effect of Macroinvertebrate Exclusion on Leaf Breakdown Rates in a Tropical Headwater Stream1

The role of macroinvertebrates in the process of leaf breakdown is well studied in temperate streams, but less is known about their role in the tropics. We investigated the effect of reducing macroinvertebrate access to leaf material on leaf breakdown rates in a forested headwater stream in the Luquillo Experimental Forest, Puerto Rico. We measured leaf mass loss using fine and coarse mesh bags over 12 weeks for two common riparian species: Cecropia schreberiana (Moraceae) and Dacryodes excelsa (Burseraceae). Coarse mesh allowed freshwater shrimp and other macroinvertebrates to access leaf material, while fine mesh did not. Leaf breakdown rates did not differ between C. schreberiana and D. excelsa in coarse mesh bags (?0.0375/day vs. ?0.0395/day, respectively), but C. schreberiana breakdown was significantly slower than D. excelsa in fine mesh bags (?0.0159/day vs. ?0.0266/day). Additionally, breakdown in fine mesh bags was significantly slower compared to coarse mesh bags for C. schreberiana, but less so for D. excelsa. Breakdown rates for all treatments were fast relative to those in temperate‐zone streams indicating that both macroinvertebrates and macroinvertebrate‐independent processing can strongly influence leaf decomposition in tropical streams. The difference between C. schreberiana and D. excelsa indicates that the effect of macroinvertebrate exclusion can change with leaf type.     

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.     

Inter-annual,Annual, and Seasonal Variation of P and N Retention in a Perennial and an Intermittent Stream

Headwater streams represent the key sites of nutrient retention, but little is known about temporal variation in this important process. We used monthly measurements over 2 years to examine variation in retention of soluble reactive phosphorus (SRP) and ammonium (NH₄⁺) in two Mediterranean headwater streams with contrasting hydrological regimes (that is, perennial versus intermittent). Differences in retention between streams were more evident for NH₄⁺, likely due to strong differences in the potential for nitrogen limitation. In both streams, nutrient-retention efficiency was negatively influenced by abrupt discharge changes, whereas gradual seasonal changes in SRP demand were partially controlled by riparian vegetation dynamics through changes in organic matter and light availability. Nutrient concentrations were below saturation in the two streams; however, SRP demand increased relative to NH₄⁺ demand in the intermittent stream as the potential for phosphorus limitation increased (that is, higher dissolved inorganic nitrogen:SRP ratio). Unexpectedly, variability in nutrient retention was not greater in the intermittent stream, suggesting high resilience of biological communities responsible for nutrient uptake. Within-stream variability of all retention metrics, however, increased with increasing time scale. A review of studies addressing temporal variation of nutrient retention at different time scales supports this finding, indicating increasing variability of nutrient retention with concomitant increases in the variability of environmental factors from the diurnal to the inter-annual scale. Overall, this study emphasizes the significance of local climate conditions in regulating nutrient retention and points to potential effects of changes in land use and climate regimes on the functioning of stream ecosystems.     

Temperature and nutrient effects on the relative importance of brown and green pathways for stream ecosystem functioning: A mesocosm approach

1. In addition to global warming, aquatic ecosystems are currently facing multiple global changes among which include changes in nitrogen (N) loads. While several studies have investigated both temperature and N impacts on aquatic ecosystems independently, knowledge on their interactive effects remains scarce.
2. In forested headwater streams, decomposition of leaf litter represents the main process ensuring the transfer of nutrients and energy to higher trophic levels, followed by autochthonous primary production, mainly ensured by phototrophic biofilms. The main aim of this study was to disentangle the independent and combined effects of temperature increase and nutrient availability on the relative importance of brown and green processes involved in stream functioning. We hypothesised that water temperature and nutrients would lead to a general increase in leaf-litter decomposition and primary production, but that the intensity of these effects would be largely modulated by competitive interactions arising between microorganisms as well as by the top-down control of microorganisms by macro-invertebrates. Macro-invertebrates would, in turn, be bottom-up controlled by microbial resources quality.
3. To test these hypotheses, we conducted a 56-day experiment in artificial streams containing leaf litter, microbial decomposers and biofilm inoculum, and an assemblage of macro-invertebrates. Two water inorganic N:phosphorus (P) ratios (33 and 100, molar ratios) and two temperatures (ambient, +2°C) were manipulated, each treatment being replicated three times. Fungal and biofilm growth as well as leaf-litter decomposition and primary production were quantified. Top-down impacts of invertebrate primary consumers on brown and green compartments were evaluated using exclosures while bottom-up control was evaluated through the measurement of resource stoichiometry and fatty acid profiles, as well as quantification of macro-invertebrate growth and survival.
4. Contrary to expectations, microbial decomposition was not significantly stimulated by nutrient or temperature manipulations, while primary production was only improved under ambient temperature. In the + 2°C treatment with high N:P, greater biofilm biomass was associated with lower fungal development, which indicates competition for nutrients in these conditions. Temperature increased macro-invertebrate growth and leaf-litter consumption, but this effect was independent of any improvement of basal resource quality, suggesting that temperature mediated changes in consumer metabolism and activity was the main mechanism involved.
5. Most of our hypotheses that were based on simplified laboratory observations have been rejected in our semi-controlled mesocosms. Our study suggests that the complexity of biological communities might greatly affect the response of ecosystems to multiple stressors, and that interactions between organisms must be explicitly taken into account when investigating the impacts of global change on ecosystem functioning.

     

Light- and nutrient-limited periphyton in low order streams of Oahu,Hawaii

To date, most studies of light- and nutrient-limited primary productivity in forested streams have been carried out in deciduous forests of temperate, continental regions. Conceptual models of light and nutrient limitation have been developed from these studies, but their restricted geographic range reduces the generality of such models. Unlike temperate continental streams, streams on tropical high islands are characterized by flashy, unpredictable discharge and riparian canopies that do not vary seasonally. These contrasting conditions suggest that patterns of light and nutrient limitation in tropical streams may differ from those in temperate streams. The effects of light, and nitrogen and phosphorus availability on periphyton accrual (measured as chlorophyll a per unit area) were investigated using field experiments in 4 low-order streams on the island of Oahu, Hawaii. Levels of chlorophyll a in partially-shaded stream pools were significantly greater than in heavily-shaded pools, and nutrient-enrichment increased the level of chlorophyll a in partially-shaded pools but not in heavily-shaded pools. In each stream, phosphate enrichment resulted in an increase in the level of chlorophyll a, but nitrate enrichment had no effect. Spates following rainstorms occur frequently in these streams, and may increase periphyton productivity by increasing the flux of nutrients to algal cells. However, differences in inorganic nitrogen and phosphorus concentrations measured during spates and baseflow were small, and during some spates, concentrations of these two nutrients declined relative to baseflow concentrations. These observations suggest that phosphorus limitation was not alleviated by spates.