Characterisation and reactivity continuum of dissolved organic matter in forested headwater catchments of Andean Patagonia

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Bacterial Growth on Photochemically Transformed Leachates from Aquatic and Terrestrial Primary Producers

Abstract We measured bacterial growth on phototransformed dissolved organic matter (DOM) leached from eight different primary producers. Leachates (10 mg C liter⁻¹) were exposed to artificial UVA + UVB radiation, or kept in darkness, for 20 h. DOM solutions were subsequently inoculated with lake water bacteria. Photoproduction of dissolved inorganic carbon (DIC), ranging from 3 to 16 μg C liter⁻¹ h⁻¹, and changes in the absorptive characteristics of the DOM were observed for all leachates upon UV irradiation. The effects of irradiation exposure on DOM bioavailability varied greatly, depending on leachate and type of bacterial growth criterion. Bacterial carbon utilization (biomass production plus respiration) over the entire incubation period (120 h) was enhanced by UV radiation of leachate from the terrestrial leaves, relative to carbon utilization in non-irradiated leachates. Conversely, carbon utilization was reduced by radiation of the leachates from aquatic macrophytes. In a separate experiment, the stable C and N isotope composition of bacteria grown on irradiated and non-irradiated DOM was estimated. Bacterial growth on UV-irradiated DOM was enriched in ¹³C relative to the bacteria in the non-irradiated treatments; this result may be explained by selective assimilation of photochemically produced, isotopically enriched labile compounds. Received: 17 February 2000; Accepted: 1 May 2000; Online Publication: 28 August 2000     

Photomineralization in a boreal hydroelectric reservoir: a comparison with natural aquatic ecosystems

In order to evaluate the role of photochemistry in the carbon dioxide (CO₂) generation from a 10-year-old boreal reservoir, the photomineralization of dissolved organic matter (DOM) was assessed and compared to a boreal river as well as to boreal and temperate lakes during July and August, 2003. Sterile water samples were irradiated by sunlight over the whole photoperiod and subsequently analyzed for CO₂. Mean energy-normalized apparent photochemical yield of CO₂ (an index of DOM photoreactivity normalized for the energy absorbed by samples) was significantly higher in the reservoir (27.7 ± 13.0 mg CO₂·m⁻³·kJ⁻¹) and the boreal river (35.8 ± 2.3 mg CO₂·m⁻³·kJ⁻¹) than in the boreal lakes (15.5 ± 5.1 mg CO₂·m⁻³·kJ⁻¹). The DOM photoreactivity of the temperate lakes (20.9 ± 8.1 mg CO₂·m⁻³·kJ⁻¹) was not statistically different from any type of boreal water bodies. There was no significant difference in either the integrated photoproduction of CO₂ (273–433 mg CO₂·m⁻²·d⁻¹) or the potential photochemical contribution to CO₂ diffusive fluxes (56–92%) among these water bodies. DOM photoreactivity was significantly affected by the cumulative hydrological residence time (CHRT) when considering the whole data set. However, when considering only the boreal water bodies, iron (Fe) and manganese (Mn) also intervened. The fact that DOM photoreactivity was related to CHRT as well as to Fe and Mn concentrations, which are respectively permanent and long-lasting features of the reservoir, suggests that the photoproduction of CO₂ is not likely to decrease over time. This process may therefore play a substantial role in the long-term CO₂ emissions from boreal reservoirs during the summer, its potential contribution to CO₂ diffusive fluxes being estimated at 56 ± 29 %.     

Effect of Wild and Genetically Modified Rhizosphere Bacteria Pseudomonas aureofaciens on the Accumulation of Arsenic by Plants

Gene constructions rendering recombinant bacteria resistant to arsenic and increasing their ability to dissolve soil phosphates and/or arsenates were created by cloning the ars operon and the gene of citrate synthase from a chromosome of the strain Pseudomonas aeruginosa PA01. Genetically modified variants of the strain Pseudomonas aureofaciens BS1393 have been constructed that are resistant to high concentrations of arsenic and dissolve poorly soluble phosphates and/or arsenates. The recombinant strains P. aureofaciens BS1393(pUCP22::arsRBC) and P. aureofaciens BS1393(pUCP22::gltA) exerted positive effects on the survival of sorgo (Sorghum saccharatum L.) and its ability to accumulate arsenic.     

Effects of citric acid and the siderophore desferrioxamine B (DFO-B) on the mobility of germanium and rare earth elements in soil and uptake in Phalaris arundinacea

Effects of citric acid and desferrioxamine B (DFO-B) on the availability of Ge and selected rare earth elements (REEs) (La, Nd, Gd, Er) to Phalaris arundinacea were investigated. A soil dissolution experiment was conducted to elucidate the effect of citric acid and DFO-B at different concentrations (1 and 10 mmol L^?1 citric acid) on the release of Ge and REEs from soil. In a greenhouse, plants of P. arundinacea were cultivated on soil and on sand cultures to investigate the effects of citric acid and DFO-B on the uptake of Ge and REEs by the plants. Addition of 10 mmol L^?1 citric acid significantly enhanced desorption of Ge and REEs from soil and uptake into soil-grown plants. Applying DFO-B enhanced the dissolution and the uptake of REEs, while no effect on Ge was observed. In sand cultures, the presence of citric acid and DFO-B significantly decreased the uptake of Ge and REEs, indicating a discrimination of the formed complexes during uptake. This study clearly indicates that citric acid and the microbial siderophore DFO-B may enhance phytoextraction of Ge and REEs due to the formation of soluble complexes that increase the migration of elements in the rhizosphere.     

The structure of a pyoverdine produced by a Pseudomonas tolaasii-like isolate

Cultures of Agaricus bisporus, the most extensively cultivated mushroom, can be infected severely by Pseudomonas tolaasii. This pathogen is characterized by the so-called white line reaction, a precipitate formed on agar plates between its colonies and those of P. reactans, both belonging to the collective species P. fluorescens. A recent study has shown that a group of P. tolaasii isolates can be subdivided into two groups or 'siderovars', based on the pyoverdines they produce (Munsch et al. 2000). One group of strains is characterized by the pyoverdine described by Demange et al. (1990). A representative of the second group (strain Ps3a) was found to produce the same pyoverdine as a strain which had been classified before as P. aureofaciens. However, based mainly on 16S rRNA gene sequence comparisons and REP-PCR generated fingerprints, the two strains are not identical. They are also distinguishable from the P. tolaasii type strain.     

Effects of macrophyte leachate on <Emphasis Type="Italic">Anabaena</Emphasis> sp. and <Emphasis Type="Italic">Chlamydomonas moewusii</Emphasis> growth in freshwater tropical ecosystems

This study reports on the effects of dissolved organic matter (DOM) derived from the aquatic macrophyte Pistia stratiotes (collected from a tropical reservoir) on the mixotrophic growth of two phytoplankton species (Chlamydomonas moewusii and Anabaena sp.). The DOM from P. stratiotes had a mainly aliphatic structure, low molecular weight, low cellulose and lignin content and high carbon content. The addition of DOM (5% v/v) significantly decreased the growth rate of Anabaena sp. but increased the chlorophyll a concentration of C. moewusii. Higher light intensity (100 versus 30 µmol m^?2 s^?1) was important for Anabaena sp., increasing its growth rate and chlorophyll content. The use of DOM from P. stratiotes to mitigate cyanobacterial blooms should be further explored in future studies.     

New record of moss and thermophilic bacteria species and physico-chemical properties of geothermal soils on the northwest slope of Mt. Melbourne (Antarctica)

Four samples of surface soils, one with shoots of an unidentified moss species, were collected from a geothermal site on the northwest slope of volcanic Mt. Melbourne (northern Victoria Land, continental Antarctica) to determine physico-chemical properties, isolate existing strains of heterotrophic microorganisms, and identify the moss species through molecular genetic techniques. Surface soil features such as temperature, grain-size, pH, moisture content, and isolated genera of bacteria, generally corresponded to those previously reported for other geothermal sites in Victoria Land. However, when compared with chemical characteristics of warm substrata from these sites, soils from the northwest slope of Mt. Melbourne showed lower contents of total N and water-extractable PO₄^3– and K⁺, and relatively higher concentrations of Na, Fe, Mn, and of potentially toxic elements such as Cd and Pb. Preliminary results indicate that a new species of thermophilic bacteria growing in Fe-enriched medium was isolated. Although the study area lay only about 1.5 km from Cryptogam Ridge, a geothermal site in the rim of the Mt. Melbourne summit crater with a well-developed population of the moss Campylopus pyriformis, molecular genetic analyses showed that the moss on the volcano slope is Pohlia nutans, a species closely related to populations some 110 km to the north in the Mt. Rittmann fumaroles. It was concluded that physico-chemical features of geothermal grounds may affect the colonisation history and dispersal of microorganisms and mosses.     

Fate of the Biological Control Agent Pseudomonas aureofaciens TX-1 after Application to Turfgrass

The fate and impact of Pseudomonas aureofaciens TX-1 following application as a biocontrol agent for fungi in turfgrass were studied. The organism was applied with a modified irrigation system by using a preparation containing 1 × 10⁶ P. aureofaciens TX-1 CFU ml⁻¹ about 100 times between May and August. We examined the impact of this repeated introduction of P. aureofaciens TX-1 (which is known to produce the antimicrobial compound phenazine-1-carboxylic acid) on the indigenous microbial community of the turfgrass system and on establishment of introduced bacteria in the soil system. A PCR primer-DNA hybridization probe combination was developed to accurately monitor the fate of P. aureofaciens TX-1 following application in irrigation water. To assess the impact of frequent P. aureofaciens TX-1 applications on the indigenous bacterial community, turfgrass canopy, thatch, and rhizosphere samples were obtained during the growing season from control and treated plots and subjected to DNA extraction procedures and denaturing gradient gel electrophoresis (DGGE). PCR amplification and hybridization of extracted DNA with the P. aureofaciens TX-1-specific primer-probe combination revealed that P. aureofaciens TX-1 not only became established in the rhizosphere and thatch but also was capable of overwintering. Separation of PCR-amplified partial 16S rRNA genes by DGGE showed that the repeated application of P. aureofaciens TX-1 in irrigation water resulted in transient displacement of a leaf surface bacterial community member. There was no obvious alteration of any dominant members of the thatch and rhizosphere microbial communities.     

Mineralisation of dissolved organic matter by heterotrophic stream biofilm communities in a large boreal catchment

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Changes in stable nitrogen and carbon isotope ratios of plants and soil across a boreal forest fire chronosequence

Background and Aim

Nitrogen (N) and carbon (C) isotopic signatures (δ¹⁵N and δ¹³C) serve as powerful tools for understanding temporal changes in ecosystem processes, but how these signatures change across boreal forest chronosequences is poorly understood.

Methods

The δ¹⁵N, δ¹³C, and C/N ratio of foliage of eight dominant plant species, including trees, understory shrubs, and a moss, as well as humus, were examined across a 361 years fire-driven chronosequence in boreal forest in northern Sweden.

Results

The δ¹³C and C/N ratio of plants and humus increased along the chronosequence, suggesting increasing plant stress through N limitation. Despite increasing biological N fixation by cyanobacteria associated with feather mosses, δ¹⁵N showed an overall decline, and δ¹⁵N of the feather moss and associated vascular plants diverged over time from that of atmospheric N₂.

Conclusions

Across this chronosequence the N fixed by cyanobacteria is unlikely to be used by mosses and vascular plants without first undergoing mineralization and mycorrhizal transport, which would cause a change in δ¹⁵N signature due to isotopic fractionation. The decreasing trend of δ¹⁵N suggests that as the chronosequence proceeds, the plants may become more dependent on N transferred from mycorrhizal fungi or from N deposition.     

Are Mosses Required to Accurately Predict Upland Black Spruce Forest Soil Carbon in National-Scale Forest C Accounting Models?

The boreal forest plays a key role in the global carbon (C) cycle, and black spruce (Picea mariana (Mill.) BSP) forests are the dominant coniferous forest type in the Canadian boreal forest. National-scale forest C models currently do not account for the contribution of moss-derived organic matter that we hypothesize to be significant in the C budget of black spruce ecosystems. One such model, the Carbon Budget Model of the Canadian Forest Sector (CBM-CFS3), is designed to meet Canada’s forest-related greenhouse gas reporting requirements. In this study our goal was to determine if black spruce forest soil C stocks are significantly underestimated by the CBM-CFS3, and if so, to determine if estimates could be improved by adding moss-derived C. We conclude that in black spruce sites, organic layer C is significantly underestimated by CBM-CFS3 compared to sites with all other leading tree species analyzed. We compiled and used published moss net primary productivity rates for upland forest systems, with decomposition rates, in mass-balance calculations to estimate mean moss-derived C in black spruce forests for feather mosses at 64 Mg C ha^?1, and for sphagnum mosses at 103 Mg C ha^?1. These C pools are similar to the CBM-CFS3 mean underestimation of black spruce soil organic layers (63 Mg C ha^?1). We conclude that the contribution of mosses is sufficiently large that a moss C pool should be added to national-scale models including the CBM-CFS3, to reduce uncertainties in boreal forest C budget estimation. Feather and sphagnum mosses should be parameterized separately.     

Nitrogen fixation in mixed <Emphasis Type="Italic">Hylocomium splendens</Emphasis> moss communities

The pleurocarpus feather moss, Hylocomium splendens, is one of two co-dominant moss species in boreal forest ecosystems and one of the most common mosses on earth, yet little is known regarding its capacity to host cyanobacterial associates and thus contribute total ecosystem N. In these studies, we evaluated the N-fixation potential of the H. splendens–cyanobacteria association and contrasted the N-fixation activity with that of the putative N-fixing moss–cyanobacteria association of Pleurozium schreberi. Studies were conducted to: quantify N-fixation in H. splendens and P. schreberi in sites ranging from southern to northern Fennoscandia; assess N and P availability as drivers of N-fixation rates; contrast season-long N-fixation rates for both mosses; and characterize the cyanobacteria that colonize shoots of H. splendens. Nitrogen-fixation rates were generally low at southern latitudes and higher at northern latitudes (64–69°N) potentially related to anthropogenic N deposition across this gradient. Nitrogen fixation in H. splendens appeared to be less sensitive to N deposition than P. schreberi. The season-long assessment of N-fixation rates at a mixed feather moss site in northern Sweden showed that H. splendens fixed a substantial quantity of N, but about 50% less total N compared to the contribution from P. schreberi. In total, both species provided 1.6 kg fixed N ha⁻¹ year⁻¹. Interestingly, H. splendens demonstrated somewhat higher N-fixation rates at high fertility sites compared to P. schreberi. Nostoc spp. and Stigonema spp. were the primary cyanobacteria found to colonize H. splendens and P. schreberi. These results suggest that H. splendens with associated Nostoc or Stigonema communities contributes a significant quantity of N to boreal forest ecosystems, but the contribution is subordinate to that of P. schreberi at northern latitudes. Epiphytic cyanobacteria are likely a key factor determining the co-dominant presence of these two feather mosses across the boreal biome.     

Mosses in High-Arctic lakes: in situ measurements of annual primary production and decomposition

Aquatic mosses are important primary producers in High-Arctic lakes, but little information is available on their contribution to the overall production in these lakes. In order to predict effects of climate change on whole-lake ecosystem characteristics, more knowledge is needed on the role of moss in primary production, the extent of nutrient limitation of moss primary production and whether moss serves as food resource for secondary producers. In this study, we conducted an in situ growth experiment of an aquatic moss in a High-Arctic lake in NE Greenland and used these data to determine annual net production of this moss in the whole lake. We also measured tissue-N and tissue-P in order to assess nutrient limitation of moss production, measured in situ decomposition rates by litter bag experiments over 1 year and assessed the role of moss as food source by analysing stable isotope ¹⁵N and ¹³C of relevant organism groups in the lake. Net primary production of moss was 1.3 gC m^?2 year^?1 and constituted 23 % of the total benthic primary production and 18 % of the total lake primary production. Stoichiometric assessments suggested N and P limitation of moss growth. On average, 15 % of the standing biomass was decomposed per year. Our results also indicate that moss is not directly used as food resource by herbivores, but the most abundant herbivore, Lepidurus arcticus, is feeding on the epiphytic biofilm on the moss. Moss biomass is instead incorporated into the microbial decomposer pathway. All together, the study shows that moss plays an important ecological role as primary producer in High-Arctic lakes and functions as substrate for periphytic biofilm that serves as food resource for important herbivore invertebrates.     

Onsite chelate resin solid-phase extraction of rare earth elements in natural water samples: its implication for studying past redox changes by inorganic geochemistry

Rare earth element (REE) patterns in natural water and geological samples provide information on previous changes in environmental conditions, such as redox changes and material cycles; however, quantitative analysis of REEs in these samples is complicated by the relatively low contents of REEs in such samples as well as mass interference from ¹³⁵Ba¹⁶O and ¹³⁷Ba¹⁶O in inductively coupled plasma mass spectrometry (ICP-MS) analyses. In this study, onsite solid-phase extraction and preconcentration methods for REEs using an iminobisacetic acid–ethylenediaminetriacetic acid chelate resin (Nobias Chelate PA1, Hitachi High-Tech Fielding) were adopted for the analyses. Standard reference materials (SPS-SW1 artificial surface water) and natural ground water and spring water samples were used to evaluate the methods. Using the chelate resin, background levels of REEs were found to be less than 0.3 ng L^?1 and recovery rates (REEs, 1 ng L^?1) were 97.9–106.7% for the artificial surface water. Ba contents were lower than the detection limit after extraction. Additionally, duplicate analyses were performed to check the reproducibility of the onsite extraction. The REE patterns in the natural water samples were in good agreement with those obtained using a previous method (the interference calibration method without solid-phase extraction). Therefore, onsite solid-phase extraction using the chelate resin was demonstrated to be a rapid and simple preparation technique for REE analyses.     

Biogenic Manganese Oxides Facilitate Iodide Oxidation at pH ≤ 5

Radioactive ¹²⁹I, a byproduct of nuclear power generation, can pose risks to human health if released into the environment, where its mobility is highly dependent on speciation. Based on thermodynamic principles, ¹²⁹I should exist primarily as iodide (I^?) in most terrestrial environments; however, organo-¹²⁹I and ¹²⁹iodate are also commonly detected in contaminated soils and groundwater. To investigate the capability of biogenic manganese oxides to influence iodide speciation, 17 manganese-oxidizing bacterial strains, representing six genera, were isolated from soils of the Savannah River Site, South Carolina. The isolates produced between 2.6 and 67.1 nmole Mn oxides (ml^?1 media after 25 days, pH 6.5). Results from inhibitor assays targeting extracellular enzymes and reactive oxygen species indicated that both play a role in microbe-induced Mn(II) oxidation among the strains examined. Iodide oxidation was not observed in cultures of the most active Mn-oxidizing bacteria, Chryseobacterium sp. strain SRS1 and Chromobacterium sp. strain SRS8, or the fungus, Acremonium strictum strain KR21–2. While substantial amounts of Mn(III/IV) oxides were only generated in cultures at ≥pH 6, iodide oxidation was only observed in the presence of Mn(III/IV) oxides when the pH was ≤5. Iodide oxidation was promoted to a greater extent by synthetic Mn(IV)O₂ than biogenic Mn(III/IV) oxides under these low pH conditions (≤pH 5). These results indicate that the influence of biogenic manganese oxides on iodide oxidation and immobilization is primarily limited to low pH environments.     

Chemical composition of aquatic dissolved organic matter in five boreal forest catchments sampled in spring and fall seasons

The chemical composition and carbon isotope signature of aquatic dissolved organic matter (DOM) in five boreal forest catchments in Scandinavia were investigated. The DOM was isolated during spring and fall seasons using a reverse osmosis technique. The DOM samples were analyzed by elemental analysis, FT-IR, solid-state CP-MAS ¹³C-NMR, and C-1s NEXAFS spectroscopy. In addition, the relative abundance of carbon isotopes (¹²C, ¹³C, ¹⁴C) in the samples was measured. There were no significant differences in the chemical composition or carbon isotope signature of the DOM sampled in spring and fall seasons. Also, differences in DOM composition between the five catchments were minor. Compared to reference peat fulvic and humic acids, all DOM samples were richer in O-alkyl carbon and contained less aromatic and phenolic carbon, as shown by FT-IR, ¹³C-NMR, and C-1s NEXAFS spectroscopy. The DOM was clearly enriched in ¹⁴C relative to the NBS oxalic acid standard of 1950, indicating that the aquatic DOM contained considerable amounts of organic carbon younger than about 50 years. The weight-based C:N ratios of 31 ± 6 and the values of indicate that the isolated DOM is of terrestrial rather than aquatic origin. We conclude that young, hydrophilic carbon compounds of terrestrial origin are predominant in the samples investigated, and that the composition of the aquatic DOM in the studied boreal forest catchments is rather stable during low to intermediate flow conditions.     

Mycorrhiza and soil bacteria influence extractable iron and manganese in soil and uptake by soybean

Excess manganese (Mn) in soil is toxic to crops, but in some situations arbuscular mycorrhizal fungi (AMF) alleviate the toxic effects of Mn. Besides the increased phosphorus (P) uptake, mycorrhiza may affect the balance between Mn-reducing and Mn-oxidizing microorganisms in the mycorrhizosphere and affect the level of extractable Mn in soil. The aim of this work was to compare mycorrhizal and non-mycorrhizal plants that received extra P in relation to alleviation of Mn toxicity and the balance between Mn-oxidizing and Mn-reducing bacteria in the mycorrhizosphere. A clayey soil containing 508 mg kg⁻¹ of extractable Mn was fertilized with 30 mg kg⁻¹ (P1) or 45 mg kg⁻¹ (P2) of soluble P. Soybean (Glycine max L. Merrill, cv. IAC 8-2) plants at P1 level were non-inoculated (CP1) or inoculated with either Glomus etunicatum (GeP1) or G. macrocarpum (GmP1), while plants at P2 level were left non-inoculated (CP2). Plants were grown in a greenhouse and harvested after 80 days. In the mycorrhizosphere of the GmP1 and GeP1 plants a shift from Mn-oxidizing to Mn-reducing bacteria coincided with higher soil extractability of Mn and Fe. However, the occurrence of Mn-oxidizing/reducing bacteria in the (mycor)rhizosphere was unrelated to Mn toxicity in plants. Using 16S rDNA sequence homologies, the Mn-reducing isolates were consistent with the genus Streptomyces. The Mn-oxidizers were homologous with the genera Arthrobacter, Variovorax and Ralstonia. While CP1 plants showed Mn toxicity throughout the whole growth period, CP2 plants never did, in spite of having Fe and Mn shoot concentrations as high as in CP1 plants. Mycorrhizal plants showed Mn toxicity symptoms early in the growth period that were no longer visible in later growth stages. The shoot P concentration was almost twice as high in mycorrhizal plants compared with CP1 and CP2 plants. The shoot Mn and Fe concentrations and contents were lower in GmP1 and GeP1 plants compared with the CP2 treatment, even though levels of extractable metals increased in the soil when plants were mycorrhizal. This suggests that mycorrhiza protected its host plant from excessive uptake of Mn and Fe. In addition, higher tissue P concentrations may have facilitated internal detoxification of Mn in mycorrhizal plants. The exact mechanisms acting on alleviation of Mn toxicity in mycorrhizal plants should be further investigated.     

Ecophysiological analysis of moss-dominated biological soil crusts and their separate components from the Succulent Karoo, South Africa

Biological soil crusts, formed by an association of soil particles with cyanobacteria, lichens, mosses, fungi and bacteria in varying proportions, live in or directly on top of the uppermost soil layer. To evaluate their role in the global carbon cycle, gas exchange measurements were conducted under controlled conditions. Moss-dominated soil crusts were first analyzed as moss tufts on soil, then the mosses were removed and the soil was analyzed separately to obtain the physiological response of both soil and individual moss stems. Net photosynthetic response of moss stems and complete crusts was decreased by insufficient and excess amounts of water, resulting in optimum curves with similar ranges of optimum water content. Light saturation of both sample types occurred at high irradiance, but moss stems reached light compensation and saturation points at lower values. Optimum temperatures of moss stems ranged between 22 and 27°C, whereas complete crusts reached similar net photosynthesis between 7 and 27°C. Under optimum conditions, moss stems reached higher net photosynthesis (4.0 vs. 2.8 μmol m^?2 s^?1) and lower dark respiration rates (?0.9 vs. ?2.4 μmol m^?2 s^?1). Respiration rates of soil without moss stems were high (up to ?2.0 μmol m^?2 s^?1) causing by far lower absolute values of NP/DR ratios of soil crusts as compared to moss stems. In carbon balances, it therefore has to be clearly distinguished between measurements of soil crust components versus complete crusts. High rates of soil respiration may be caused by leaching of mosses, creating high-nutrient microsites that favor microorganism growth.     

Coupling dissolved organic carbon,CO2 and productivity in boreal lakes

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