Phosphorus and DOC availability influence the partitioning between bacterioplankton production and respiration in tidal marsh ecosystems

The organic carbon consumed by aquatic bacteria (BCC) is partitioned between bacterial production (BP) and respiration (BR), but the factors that determine BCC and its partition into BP and BR are not well understood. We explored the coupling between BR, BR and BCC, and their links to dissolved organic carbon (DOC) and nutrient availability in natural and restored tidal marshes and in the adjoining waters of Delaware Bay estuary. Labile DOC (LDOC) ranged from 3% to 22% of the DOC pool, and explained more of the variance in both BR and BCC than did bulk DOC. Bacterial growth efficiency (BGE) was highly variable (0.09-0.58), and natural Spartina alterniflora marshes had consistently higher BGE than both restoration marshes and tidal floodwaters. BGE was negatively related to the ratio of LDOC to total dissolved phosphorous, which was highest in natural marshes. The enhancement of BP observed in the marshes relative to the estuarine floodwaters had different origins: In natural marshes it was mostly due to increases in BGE, whereas in restored marshes it followed increased BCC. These results highlight the importance of P in regulating microbial metabolism in coastal areas, and the need to understand the pathways that lead to BP in these systems.     

Effects of terrigenous organic substrates and additional phosphorus on bacterioplankton metabolism and exoenzyme stoichiometry

1. Bamboo, as a pioneer vegetation, often forms forests on bare lands after catastrophic landslides. Compared to evergreen forest soil, bamboo forest soil is much more labile, with a higher percentage of microbially derived organic carbon (OC), lower molecular weight, and lower humic acid content. We hypothesised that different terrigenous organic matter (tOM) sources with varying lability and phosphorus (P) availability select for bacterioplankton with distinct metabolic pathways.
2. We incubated natural bacterioplankton assemblages with tOM leached from bamboo forest soil (BOM) and evergreen forest soil (EOM) and compared these to a lake water control. To test if microbial metabolism would be limited by OC or P availability of each tOM treatment, we used acetate as an extra labile OC source and phosphate as an inorganic P source. Bacterial metabolism was measured by analysing respiration via O₂ consumption and production via tritiated thymidine (TdR) assimilation.
3. Bacterioplankton metabolism is limited by the availability of P in BOM substrates. When using BOM, bacteria had higher enzymatic activities for phosphatase. The nutrients required for bacterial biomass seemed to be derived from organic matter. Under BOM treatment, bacterial production (BP) (0.92 ± 0.13 μg C L⁻¹ hr⁻¹) and cell specific TdR assimilation rates (0.015 ± 0.002 10^–18 M TdR cell⁻¹ hr⁻¹) were low. Adding P enhanced BP (BOM_+P 1.52 ± 0.31 and BOM_+C+P 2.25 ± 0.37 μg C L⁻¹ hr⁻¹) while acetate addition had no significant effect on BOM treatment.
4. This indicated that the bacteria switched to using added inorganic P to respire a P-limited BOM substrate, which increased total BP and abundance, resulting in even more active respiration and lower growth efficiency. We also found higher activities for chitin-degrading enzyme β-N-acetylglucosaminidase, which is associated with N mining from aminosaccharides.
5. Microbes using EOM, however, did not change metabolic strategies with additional acetate or/and inorganic P. This is due to higher concentrations of organic P in EOM substrates and the presence of inorganic N in the EOM leachates an alternative nutrient source. Bacteria produced β-glucosidase and leucyl-aminopeptidase in order to utilise the humic substances, which sustained greater bacterial abundance, higher BP (2.64 ± 0.39 μg C L⁻¹ hr⁻¹), and lower cell-specific respiration. This yielded a much higher bacterial growth efficiency (15 ± 9.2%) than the lake water control.
6. Our study demonstrated the aquatic metabolic discrepancy between tOM of different forest types. Bacterioplankton in BOM and EOM exhibit distinct metabolic responses. Bacterial metabolic strategy when using BOM implied that the supposedly stabilised biomass OM might be efficiently used by aquatic bacterioplankton. As the labile and nutrient-deficient BOM is more susceptible to the influence of additional nutrients, fertiliser residues in bamboo forest catchments might have a stronger effect on aquatic bacterial metabolic pathways. Thus, it is important to take tOM differences into consideration when building models to estimate soil carbon turnover rates along a terrestrial–aquatic continuum.

     

Bacterial Growth Efficiency in a Tropical Estuary: Seasonal Variability Subsidized by Allochthonous Carbon

Bacterial growth efficiency (BGE) is a key factor in understanding bacterial influence on carbon flow in aquatic ecosystems. We report intra-annual variability in BGE, and bacteria-mediated carbon flow in the tropical Mandovi and Zuari estuaries (southwest India) and the adjoining coastal waters (Arabian Sea). BGE ranged from 3% to 61% and showed clear temporal variability with significantly (ANOVA, p < 0.01) higher values in the estuaries (mean, 28 ± 14%) than coastal waters (mean, 12 ± 6%). The greater variability of BGE in the estuaries than coastal waters suggest some systematic response to nutrient composition and the variability of dissolved organic matter pools, as BGE was governed by bacterial secondary production (BP). Monsoonal rains and its accompanied changes brought significant variability in BGE and bacterial productivity/primary productivity (BP/PP) ratio when compared to nonmonsoon seasons in the estuaries and coastal waters. High BP/PP ratio (>1) together with high carbon flux through bacteria (>100% of primary productivity) in the estuarine and coastal waters suggests that bacterioplankton consumed dissolved organic carbon in excess of the amount produced in situ by phytoplankton of this region, which led to the mismatch between primary production of carbon and amount of carbon consumed by bacteria. Despite the two systems being subsidized by allochthonous inputs, the low BGE in the coastal waters may be attributable to the nature and time interval in the supply of allochthonous carbon.     

Nutrient availability affects the response of the calcifying chlorophyte Halimeda opuntia (L.) J.V. Lamouroux to low pH

Atmospheric carbon dioxide emissions cause a decrease in the pH and aragonite saturation state of surface ocean water. As a result, calcifying organisms are expected to suffer under future ocean conditions, but their physiological responses may depend on their nutrient status. Because many coral reefs experience high inorganic nutrient loads or seasonal changes in nutrient availability, reef organisms in localized areas will have to cope with elevated carbon dioxide and changes in inorganic nutrients. Halimeda opuntia is a dominant calcifying primary producer on coral reefs that contributes to coral reef accretion. Therefore, we investigated the carbon and nutrient balance of H. opuntia exposed to elevated carbon dioxide and inorganic nutrients. We measured tissue nitrogen, phosphorus and carbon content as well as the activity of enzymes involved in inorganic carbon uptake and nitrogen assimilation (external carbonic anhydrase and nitrate reductase, respectively). Inorganic carbon content was lower in algae exposed to high CO₂, but calcification rates were not significantly affected by CO₂ or inorganic nutrients. Organic carbon was positively correlated to external carbonic anhydrase activity, while inorganic carbon showed the opposite correlation. Carbon dioxide had a significant effect on tissue nitrogen and organic carbon content, while inorganic nutrients affected tissue phosphorus and N:P ratios. Nitrate reductase activity was highest in algae grown under elevated CO₂ and inorganic nutrient conditions and lowest when phosphate was limiting. In general, we found that enzymatic responses were strongly influenced by nutrient availability, indicating its important role in dictating the local responses of the calcifying primary producer H. opuntia to ocean acidification.     

Short-term effects of phosphorus addition and pH rise on bacterial utilization and biodegradation of dissolved organic carbon (DOC) from boreal mires

Natural mires and forested peatlands are known to be very significant sources of dissolved organic carbon (DOC) to aquatic ecosystems. Peatland management operations (e.g., forestry operations, restoration of drained mires and peat mining) and extreme hydrological events may increase the DOC runoff. We hypothesized that an increase in phosphorus (P) leaching, together with near-neutral conditions in recipient lakes will accelerate decomposition of DOC that originates from acidic, nutrient-poor mire waters. The efficiency of DOC utilization was evaluated by measuring microbial respiration and bacterial production (BP) in short-term laboratory experiments with runoff waters from six boreal mire sites. Mere inorganic phosphorus (PO₄-P) addition did not affect the rate of respiration or the proportion of decayed DOC. However, in the nutrient-poor bog waters, P addition slightly promoted BP and bacterial growth efficiency (BGE). In contrast, the elevation of pH alone, and the elevation of pH and PO₄-P level together, caused a significant increase in respiration and in the proportion of decayed DOC, but did not affect net BP. Elevated pH alone, however, depressed BGE when compared to that under the combined elevation of pH and PO₄-P. These results suggest that the increased P availability, e.g., after mire restoration, would slightly benefit bacterial net growth in P-limited waters. However, in near-neutral recipient lakes, the increased microbial decomposition of mire-originated DOC contributes more to carbon dioxide (CO₂) supersaturation than potentially supporting detritus-based food chains.     

Relationship between Bacterioplankton Richness, Respiration, and Production in the Southern North Sea

We investigated the relationship between bacterioplankton production (BP), respiration (BR), and community composition measured by terminal restriction fragment length polymorphism in the southern North Sea over a seasonal cycle. Major changes in bacterioplankton richness were apparent from April to December. While cell-specific BP decreased highly significantly with increasing bacterioplankton richness, cell-specific BR was found to be variable along the richness gradient, suggesting that bacterioplankton respiration is rather independent from shifts in the bacterial community composition. As a consequence, the bacterial growth efficiency [BGE = BP/(BP + BR)] was negatively related to bacterioplankton richness, explaining ~43% of the variation in BGE. Our results indicate that despite the observed shifts in the community composition, the main function of the bacterioplankton, the remineralization of dissolved organic carbon to CO₂, is rather stable.     

CO2 Saturation and Trophic Shift Induced by Microbial Metabolic Processes in a River-Dominated Ocean Margin (Tropical Shallow Lagoon,Chilika, India)

An effort has been made for the first time in Asia's largest brackish water lagoon, Chilika, to investigate the spatio-temporal variability in primary productivity (PP), bacterial productivity (BP), bacterial abundance (BA), bacterial respiration (BR) and bacterial growth efficiency (BGE) in relation to partial pressure of CO₂ (pCO₂) and CO₂ air–water flux and the resultant trophic switchover. Annually, PP ranged between 24 and 376 µg C L^?1 d^?1 with significantly low values throughout the monsoon (MN), caused by light limitation due to inputs of riverine suspended matter. On the contrary, BP and BR ranged from 11.5 to 186.3 µg C L^?1 d^?1 and from 14.1 to 389.4 µg C L^?1 d^?1, respectively, with exceptionally higher values during MN. A wide spatial and temporal variation in the lagoon trophic status was apparent from BP/PP (0.05–6.4) and PP/BR (0.10–18.2) ratios. The seasonal shift in net pelagic production from autotrophy to heterotrophy due to terrestrial organic matter inputs via rivers, enhanced the bacterial metabolism during the MN, as evident from the high pCO₂ (10,134 µatm) and CO₂ air–water flux (714 mm m^?2 d^?1). Large variability in BGE and BP/PP ratios especially during MN led to high bacteria-mediated carbon fluxes which was evident from significantly high bacterial carbon demand (BCD >100% of PP) during this season. This suggested that the net amount of organic carbon (either dissolved or particulate form) synthesized by primary producers in the lagoon was not sufficient to satisfy the bacterial carbon requirements. Lagoon sustained low to moderate autotrophic–heterotrophic coupling with annual mean BCD of 231% relative to the primary production, which depicted that bacterioplankton are the mainstay of the lagoon biogeochemical cycles and principal players that bring changes in trophic status. Study disclosed that the high CO₂ supersaturation and oxygen undersaturation during MN was attributed to the increased heterotrophic respiration (in excess of PP) fuelled by allochthonous organic matter. On a spatial scale, lagoon sectors such as south sector, central sector and outer channel recorded “net autotrophic,” while the northern sector showed “net heterotrophic” throughout the study period.     

Respiration and bacterial carbon dynamics in Arctic sea ice

Bacterial carbon demand, an important component of ecosystem dynamics in polar waters and sea ice, is a function of both bacterial production (BP) and respiration (BR). BP has been found to be generally higher in sea ice than underlying waters, but rates of BR and bacterial growth efficiency (BGE) are poorly characterized in sea ice. Using melted ice core incubations, community respiration (CR), BP, and bacterial abundance (BA) were studied in sea ice and at the ice–water interface (IWI) in the Western Canadian Arctic during the spring and summer 2008. CR was converted to BR empirically. BP increased over the season and was on average 22 times higher in sea ice as compared with the IWI. Rates in ice samples were highly variable ranging from 0.2 to 18.3 μg C l⁻¹ d⁻¹. BR was also higher in ice and on average ~10 times higher than BP but was less variable ranging from 2.39 to 22.5 μg C l⁻¹ d⁻¹. Given the high variability in BP and the relatively more stable rates of BR, BP was the main driver of estimated BGE (r ² = 0.97, P < 0.0001). We conclude that microbial respiration can consume a significant proportion of primary production in sea ice and may play an important role in biogenic CO₂ fluxes between the sea ice and atmosphere.     

Temperature and the metabolic balance of streams

1. It is becoming increasingly clear that fresh waters play a major role in the global C cycle. Stream ecosystem respiration (ER) and gross primary productivity (GPP) exert a significant control on organic carbon fluxes in fluvial networks. However, little is known about how climate change will influence these fluxes. 2. Here, we used a ‘natural experiment’ to demonstrate the role of temperature and nutrient cycling in whole‐system metabolism (ER, GPP and net ecosystem production – NEP), in naturally heated geothermal (5–25 °C) Icelandic streams. 3. We calculated ER and GPP with a new, more accurate method, which enabled us to take into account the additional uncertainties owing to stream spatial heterogeneity in oxygen concentrations within a reach. ER ranged 1–25 g C m^?2 day^?1 and GPP 1–10 g C m^?2 day^?1. The median uncertainties (based on 1 SD) in ER and GPP were 50% and 20%, respectively. 4. Despite extremely low water nutrient concentrations, high metabolic rates in the warm streams were supported by fast cycling rates of nutrients, as revealed from inorganic nutrient (N, P) addition experiments. 5. ER exceeded GPP in all streams (with average GPP/ER = 0.6) and was more strongly related to temperature than GPP, resulting in elevated negative NEP with warming. We show that, as a first approximation based on summer investigations, global stream carbon emission to the atmosphere would nearly double from 0.12 Pg C year^?1 at 13 °C to 0.21 (0.15–0.33) Pg C year^?1 with a 5 °C warming. 6. Compared to previous studies from natural systems (including terrestrial ecosystems), the temperature dependence of stream metabolism was not confounded by latitude or altitude, seasonality, light and nutrient availability, water chemistry, space availability (water transient storage), and water availability. 7. Consequently, stream nutrient processing is likely to increase with warming, protecting downstream ecosystems (rivers, estuaries, coastal marine systems) during the summer low flows from nutrient enrichment, but at the cost of increased CO₂ flux back to the atmosphere.     

Bacterioplankton Growth and Nutrient Use Efficiencies Under Variable Organic Carbon and Inorganic Phosphorus Ratios

We carried out enclosure experiments in an unproductive lake in northern Sweden and studied the effects of enrichment with different dissolved organic carbon (glucose)/inorganic phosphorous (DOC/Pi) ratios on bacterioplankton production (BP), growth efficiency (BGE), nutrient use efficiency (BNUE), growth rate, and specific respiration. We found considerable variation in BP, BGE, and BNUE along the tested DOC/Pi gradient. BGE varied between 0.87 and 0.24, with the highest values at low DOC/Pi ratios. BNUE varied between 40 and 9 g C g P⁻¹, with high values at high DOC/Pi ratios. More DOC was thus allocated to growth when bacteria tended to be C-limited, and to respiration when bacteria were P-limited. Specific respiration was positively correlated with bacterial growth rate throughout the gradient. It is therefore possible that respiration was used to support growth in P-limited bacteria. The results indicated that BP can be limited by Pi when BNUE is at its maximum, by organic C when BGE is at its maximum, and by dual organic C and Pi limitation when BNUE and BGE have suboptimal values.     

Establishment of Microbial Eukaryotic Enrichment Cultures from a Chemically Stratified Antarctic Lake and Assessment of Carbon Fixation Potential

Lake Bonney is one of numerous permanently ice-covered lakes located in the McMurdo Dry Valleys, Antarctica. The perennial ice cover maintains a chemically stratified water column and unlike other inland bodies of water, largely prevents external input of carbon and nutrients from streams. Biota are exposed to numerous environmental stresses, including year-round severe nutrient deficiency, low temperatures, extreme shade, hypersalinity, and 24-hour darkness during the winter ¹. These extreme environmental conditions limit the biota in Lake Bonney almost exclusively to microorganisms ².Single-celled microbial eukaryotes (called "protists") are important players in global biogeochemical cycling ³ and play important ecological roles in the cycling of carbon in the dry valley lakes, occupying both primary and tertiary roles in the aquatic food web. In the dry valley aquatic food web, protists that fix inorganic carbon (autotrophy) are the major producers of organic carbon for organotrophic organisms ^{4, 2}. Phagotrophic or heterotrophic protists capable of ingesting bacteria and smaller protists act as the top predators in the food web ⁵. Last, an unknown proportion of the protist population is capable of combined mixotrophic metabolism ^{6, 7}. Mixotrophy in protists involves the ability to combine photosynthetic capability with phagotrophic ingestion of prey microorganisms. This form of mixotrophy differs from mixotrophic metabolism in bacterial species, which generally involves uptake dissolved carbon molecules. There are currently very few protist isolates from permanently ice-capped polar lakes, and studies of protist diversity and ecology in this extreme environment have been limited ^{8, 4, 9, 10, 5}. A better understanding of protist metabolic versatility in the simple dry valley lake food web will aid in the development of models for the role of protists in the global carbon cycle.We employed an enrichment culture approach to isolate potentially phototrophic and mixotrophic protists from Lake Bonney. Sampling depths in the water column were chosen based on the location of primary production maxima and protist phylogenetic diversity ^{4, 11}, as well as variability in major abiotic factors affecting protist trophic modes: shallow sampling depths are limited for major nutrients, while deeper sampling depths are limited by light availability. In addition, lake water samples were supplemented with multiple types of growth media to promote the growth of a variety of phototrophic organisms.RubisCO catalyzes the rate limiting step in the Calvin Benson Bassham (CBB) cycle, the major pathway by which autotrophic organisms fix inorganic carbon and provide organic carbon for higher trophic levels in aquatic and terrestrial food webs ¹². In this study, we applied a radioisotope assay modified for filtered samples ¹³ to monitor maximum carboxylase activity as a proxy for carbon fixation potential and metabolic versatility in the Lake Bonney enrichment cultures.     

Influence of dissolved organic matter and inorganic nutrients on the biofilm bacterial community on artificial substrates in a northeastern Ohio, USA, stream

Stream bacteria may be influenced by the composition and availability of dissolved organic matter (DOM) and inorganic nutrients, but knowledge about how individual phylogenetic groups in biofilm are affected is still limited. In this study, the influence of DOM and inorganic nutrients on stream biofilm bacteria was examined. Biofilms were developed on artificial substrates (unglazed ceramic tiles) for 21 days in a northeastern Ohio (USA) stream for five consecutive seasons. Then, the developed biofilm assemblages were exposed, in the laboratory, to DOM (glucose, leaf leachate, and algal exudates) and inorganic nutrients (nitrate, phosphate, and nitrate and phosphate in combination) amendments for 6 days. Bacterial numbers in the biofilms were generally higher in response to the DOM treatments than to the inorganic nutrient treatments. There were also apparent seasonal variations in the response patterns of the individual bacterial taxa to the nutrient treatments; an indication that limiting resources to bacteria in stream biofilms may change over time. Overall, in contrast to the other treatments, bacterial abundance was generally highest in response to the low-molecular-weight DOM (i.e., glucose) treatment. These results further suggest that there are interactions among the different bacterial groups in biofilms that are impacted by the associated nutrient dynamics among seasons in stream ecosystems.     

Nutrients and Other Abiotic Factors Affecting Bacterial Communities in an Ohio River (USA)

Nitrogen and phosphorus additions from anthropogenic sources can alter the nutrient pool of aquatic systems, both through increased nutrient concentrations and changes in stoichiometry. Because bacteria are important in nutrient cycling and aquatic food webs, information about how nutrients affect bacterial communities enhances our understanding of how changes in nutrient concentrations and stoichiometry potentially affect aquatic ecosystems as a whole. In this study, bacterial communities were examined in biofilms from cobbles collected across seasons at three sites along the Mahoning River (Ohio) with differing levels of inorganic nutrient inputs. Members of the alpha-, beta-, and gamma-proteobacteria, the Cytophaga–Flavobacteria cluster, and the Domain Bacteria were enumerated using fluorescent in situ hybridization. Detrended canonical correspondence analysis (DCCA) revealed that stoichiometric ratios, especially the dissolved inorganic nitrogen (DIN):soluble reactive phosphorus (SRP) molar ratio (NO₂/NO₃ + NH₄:soluble reactive phosphorus), were correlated with abundance of the various bacterial taxa. However, the patterns were complicated by correlations with single nutrient concentrations and seasonal changes in temperature. Seasonal cycles appeared to play an important role in structuring the community, as there were distinct winter communities and temperature was negatively correlated with abundance of both alpha-proteobacteria and Cytophaga–Flavobacteria. However, nutrients and stoichiometry also appeared to affect the community. Numbers of cells hybridizing the Domain Bacteria probe were correlated with the DOC:DIN ratio, the beta-proteobacteria had a negative correlation with soluble reactive phosphorus concentrations and a positive correlation with the DIN:SRP ratio, and the Cytophaga–Flavobacteria had a significant negative partial correlation with the DIN:SRP ratio. Abundances of the alpha- or gamma-proteobacteria were not directly correlated to nutrient concentrations or stoichiometry. It appears that nutrient stoichiometry may be an important factor structuring bacterial communities; however, it is one of many factors, such as temperature, that are interlinked and must be considered together when studying environmental bacteria.     

Temperature and precipitation drive temporal variability in aquatic carbon and GHG concentrations and fluxes in a peatland catchment

The aquatic pathway is increasingly being recognized as an important component of catchment carbon and greenhouse gas (GHG) budgets, particularly in peatland systems due to their large carbon store and strong hydrological connectivity. In this study, we present a complete 5‐year data set of all aquatic carbon and GHG species from an ombrotrophic Scottish peatland. Measured species include particulate and dissolved forms of organic carbon (POC, DOC), dissolved inorganic carbon (DIC), CO₂, CH₄ and N₂O. We show that short‐term variability in concentrations exists across all species and this is strongly linked to discharge. Seasonal cyclicity was only evident in DOC, CO₂ and CH₄ concentration; however, temperature correlated with monthly means in all species except DIC. Although the temperature correlation with monthly DOC and POC concentrations appeared to be related to biological productivity in the terrestrial system, we suggest the temperature correlation with CO₂ and CH₄ was primarily due to in‐stream temperature‐dependent solubility. Interannual variability in total aquatic carbon concentration was strongly correlated with catchment gross primary productivity (GPP) indicating a strong potential terrestrial aquatic linkage. DOC represented the largest aquatic carbon flux term (19.3 ± 4.59 g C m^?2 yr^?1), followed by CO₂ evasion (10.0 g C m^?2 yr^?1). Despite an estimated contribution to the total aquatic carbon flux of between 8 and 48%, evasion estimates had the greatest uncertainty. Interannual variability in total aquatic carbon export was low in comparison with variability in terrestrial biosphere–atmosphere exchange, and could be explained primarily by temperature and precipitation. Our results therefore suggest that climatic change is likely to have a significant impact on annual carbon losses through the aquatic pathway, and as such, aquatic exports are fundamental to the understanding of whole catchment responses to climate change.     

Effects of light, dissolved organic carbon, and inorganic nutrients [2pt] on the relationship between algae and heterotrophic bacteria in stream periphyton

Depending on the chemical and physical environment, algae and heterotrophic bacteria in stream periphyton communities likely engage in both positive and negative interactions. We tested the hypothesis that bacteria are more closely associated with algae when allochthonous sources of labile DOC are low and algae are not light limited. Secondly, we tested the hypothesis that, under extremely oligotrophic conditions, bacteria will out-compete algae for inorganic nutrients if their carbon requirements are met by allochthonous sources. Experiments were carried out using in situ light manipulations and nutrient diffusing substrates (releasing inorganic nutrients and /or glucose) in Harts Run, an oligotrophic stream located in north central Kentucky. Although we found that both algal and bacterial biomass were higher under ambient light, bacteria did not respond to glucose in the dark. This may indicate that bacteria were associated with algae not as a carbon source, but as a substrate for colonization. In the nutrient × glucose manipulation, we found that bacteria were co-limited by inorganic nutrients. There was no evidence of algae being negatively affected by competition with bacteria for nitrogen and phosphorus. Although low temperatures might have played a role in preventing inorganic nutrient competition between these two groups of organisms, the results of both experiments may indicate that the quantitative link between periphytic bacteria and algae is stronger under oligotrophic conditions.     

Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water

The transition zone between terrestrial and freshwater habitats is highly dynamic, with large variability in environmental characteristics. Here, we investigate how these characteristics influence the nutritional status and performance of plant life forms inhabiting this zone. Specifically, we hypothesised that: (i) tissue nutrient content differs among submerged, amphibious and terrestrial species, with higher content in submerged species; and (ii) PNUE gradually increases from submerged over amphibious to terrestrial species, reflecting differences in the availability of N and P relative to inorganic C across the land–water ecotone. We found that tissue nutrient content was generally higher in submerged species and C:N and C:P ratios indicated that content was limiting for growth for ca. 20% of plant individuals, particularly those belonging to amphibious and terrestrial species groups. As predicted, the PNUE increased from submerged over amphibious to terrestrial species. We suggest that this pattern reflects that amphibious and terrestrial species allocate proportionally more nutrients into processes of importance for photosynthesis at saturating CO₂ availability, i.e. enzymes involved in substrate regeneration, compared to submerged species that are acclimated to lower availability of CO₂ in the aquatic environment. Our results indicate that enhanced nutrient loading may affect relative abundance of the three species groups in the land–water ecotone of stream ecosystems. Thus, species of amphibious and terrestrial species groups are likely to benefit more from enhanced nutrient availability in terms of faster growth compared to aquatic species, and that this can be detrimental to aquatic species growing in the land–water ecotone, e.g. Ranunculus and Callitriche.     

Seasonal Response of Stream Biofilm Communities to Dissolved Organic Matter and Nutrient Enrichments

Dissolved organic matter (DOM) and inorganic nutrients may affect microbial communities in streams, but little is known about the impact of these factors on specific taxa within bacterial assemblages in biofilms. In this study, nutrient diffusing artificial substrates were used to examine bacterial responses to DOM (i.e., glucose, leaf leachate, and algal exudates) and inorganic nutrients (nitrate and phosphate singly and in combination). Artificial substrates were deployed for five seasons, from summer 2002 to summer 2003, in a northeastern Ohio stream. Differences were observed in the responses of bacterial taxa examined to various DOM and inorganic nutrient treatments, and the response patterns varied seasonally, indicating that resources that limit the bacterial communities change over time. Overall, the greatest responses were to labile, low-molecular-weight DOM (i.e., glucose) at times when chlorophyll a concentrations were low due to scouring during significant storm events. Different types of DOM and inorganic nutrients induced various responses among bacterial taxa in the biofilms examined, and these responses would not have been apparent if they were examined at the community level or if seasonal changes were not taken into account.     

Distinct effects of N and P addition on soil enzyme activities and C distribution in aggregates in a subalpine spruce plantation

We assessed stream ecosystem-level response to a drought-induced defoliation event by gypsy moth caterpillars (Lymantria dispar) with high-frequency water quality sensors. The defoliation event was compared to the prior year of data. Based on long-term records of precipitation and drought indices, the drought of 2015–2016 in Rhode Island, USA was an extreme climatic event that preceded and likely precipitated the defoliation from insect infestation. Canopy cover in the riparian area was reduced by over 50% increasing light availability which warmed the stream and stimulated autotrophic activity. Frass and leaf detritus contributed particulate carbon and organic nutrients to the stream. Based on locally calibrated s::can spectro::lyser data, nitrate concentration and flux did not significantly increase during defoliation while orthophosphate concentration and flux did significantly increase during part of the defoliation period. Lower mean daily dissolved oxygen (DO) levels and wider diel cycles of DO indicated higher biological activity during the defoliation event. Stream metabolism metrics were also significantly higher during defoliation and pointed to heterotrophic activity dominating in the stream. The increases in stream metabolism were low compared to other studies; in streams with higher nutrient levels (e.g., in agricultural or urban watersheds) the increase in light and temperature could have a stronger influence on stream metabolism. The in-stream metabolic processes and nutrient fluxes observed in response to the drought-driven defoliation event resulted from the long-term deployment of high-frequency water sensors. The proliferation of these water sensors now enable studies that assess ecosystem responses to stochastic, unusual disturbances.     

Large predators and biogeochemical hotspots: brown bear (Ursus arctos) predation on salmon alters nitrogen cycling in riparian soils

Two important themes in ecology include the understanding of how interactions among species control ecosystem processes, and how habitats can be connected through transfers of nutrients and energy by mobile organisms. An impressive example of both is the large influx of nutrients and organic matter that anadromous salmon supply to inland aquatic and terrestrial ecosystems and the role of predation by brown bears (Ursus arctos) in transferring these marine-derived nutrients (MDN) from freshwater to riparian habitats. In spite of the recognition that salmon-bear interactions likely play an important role in controlling the flux of MDN from aquatic to riparian habitats, few studies have linked bear predation on salmon to processes such as nitrogen (N) or carbon (C) cycling. We combine landscape-level survey data and a replicated bear-exclosure experiment to test how bear foraging on salmon affects nitrous oxide (N₂O) flux, carbon dioxide (CO₂) flux, and nutrient concentrations of riparian soils. Our results show that bears feeding on salmon increased soil ammonium (NH₄ ⁺) concentrations three-fold and N₂O flux by 32-fold. Soil CO₂ flux, nitrate (NO₃ ⁻), and N transformation differences were negligible in areas where bears fed on salmon. Reference areas without concentrated bear activity showed no detectable change in soil N cycling after the arrival of salmon to streams. Exclosure experiments showed that bear effects on soil nutrient cycles were transient, and soil N processing returned to background conditions within 1 year after bears were removed from the system. These results suggest that recipient ecosystems do not show uniform responses to MDN inputs and highlight the importance of large mobile consumers in generating landscape heterogeneity in nutrient cycles.     

Nitrogen limitation of heterotrophic biofilms in boreal streams

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