Legacy effects of drought alters the aquatic food web of a northern boreal peatland

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Rapid Response of Hydrological Loss of DOC to Water Table Drawdown and Warming in Zoige Peatland: Results from a Mesocosm Experiment

A large portion of the global carbon pool is stored in peatlands, which are sensitive to a changing environment conditions. The hydrological loss of dissolved organic carbon (DOC) is believed to play a key role in determining the carbon balance in peatlands. Zoige peatland, the largest peat store in China, is experiencing climatic warming and drying as well as experiencing severe artificial drainage. Using a fully crossed factorial design, we experimentally manipulated temperature and controlled the water tables in large mesocosms containing intact peat monoliths. Specifically, we determined the impact of warming and water table position on the hydrological loss of DOC, the exported amounts, concentrations and qualities of DOC, and the discharge volume in Zoige peatland. Our results revealed that of the water table position had a greater impact on DOC export than the warming treatment, which showed no interactive effects with the water table treatment. Both DOC concentration and discharge volume were significantly increased when water table drawdown, while only the DOC concentration was significantly promoted by warming treatment. Annual DOC export was increased by 69% and 102% when the water table, controlled at 0 cm, was experimentally lowered by −10 cm and −20 cm. Increases in colored and aromatic constituents of DOC (measured by Abs_{254 nm}, SUVA_{254 nm}, Abs_{400 nm}, and SUVA_{400 nm}) were observed under the lower water tables and at the higher peat temperature. Our results provide an indication of the potential impacts of climatic change and anthropogenic drainage on the carbon cycle and/or water storage in a peatland and simultaneously imply the likelihood of potential damage to downstream ecosystems. Furthermore, our results highlight the need for local protection and sustainable development, as well as suggest that more research is required to better understand the impacts of climatic change and artificial disturbances on peatland degradation.     

Warming enhances the stimulatory effect of algal exudates on dissolved organic carbon decomposition

1. The current paradigm in peatland ecology is that the organic matter inputs from plant photosynthesis (e.g. moss litter) exceed that of decomposition, tipping the metabolic balance in favour of carbon (C) storage. Here, we investigated an alternative hypothesis, whereby exudates released by microalgae can actually accelerate C losses from the surface waters of northern peatlands by stimulating dissolved organic C (DOC) decomposition in a warmer environment expected with climate change. To test this hypothesis, we evaluated the biodegradability of fen DOC in a factorial design with and without algal DOC in both ambient (15°C) and elevated (20°C) water temperatures during a laboratory bioassay.
2. When DOC sources were evaluated separately, decomposition rates were higher in treatments with algal DOC only than with fen DOC only, indicating that the quality of the organic matter influenced degradability. A mixture of substrates (½ algal DOC + ½ fen DOC) exceeded the expected level of biodegradation (i.e. the average of the individual substrate responses) by as much as 10%, and the magnitude of this effect increased to more than 15% with warming.
3. Specific ultraviolet absorbance at 254 nm (SUVA₂₅₄), a proxy for aromatic content, was also significantly higher (i.e. more humic) in the mixture treatment than expected from SUVA₂₅₄ values in single substrate treatments.
4. Accelerated decomposition in the presence of algal DOC was coupled with an increase in bacterial biomass, demonstrating that enhanced metabolism was associated with a more abundant microbial community.
5. These results present an alternative energy pathway for heterotrophic consumers to breakdown organic matter in northern peatlands. Since decomposition in northern peatlands is often limited by the availability of labile organic matter, this mechanism could become increasingly important as a pathway for decomposition in the surface waters of northern peatlands where algae are expected to be more abundant in conditions associated with ongoing climate change.

     

Global warming and the export of dissolved organic carbon from boreal peatlands

Peatlands occupy approximately 15% of boreal and sub-arctic regions, contain approximately one third of the world's soil carbon pool, and supply most of the dissolved organic carbon (DOC) entering boreal lakes and rivers and the Arctic Ocean. The high latitudes occupied by these peatlands are expected to see the greatest amount of climatic warming in the next several decades. In addition to increasing temperatures, climatic change could also affect the position of the water-table level and discharge from these peatlands. Changes in temperature, water tables, and discharge could affect delivery of DOC to downstream ecosystems where it exerts significant control over productivity, biogeochemical cycles, and attenuation of visible and UV radiation. We experimentally warmed and controlled water tables while measuring discharge in a factorial experiment in large mesocosms containing peat monoliths and intact plant communities from a bog and fen to determine the effects of climate change on DOC budgets. We show that the DOC budget is controlled largely by changes in discharge rather than by any effect of warming or position of the water-table level on DOC concentrations. Furthermore, we identify a critical discharge rate in bogs and fens for which the DOC budget switches from net export to net retention. We also demonstrate an exponential increase in trace gas CO₂–C and CH₄–C emissions coincident with increased retention of dissolved organic carbon from boreal peatlands.     

Increased temperature sensitivity of net DOC production from ombrotrophic peat due to water table draw-down

The production and release of dissolved organic carbon (DOC) from peat soils is thought to be sensitive to changes in climate, specifically changes in temperature and rainfall. However, little is known about the actual rates of net DOC production in response to temperature and water table draw‐down, particularly in comparison to carbon dioxide (CO₂) fluxes. To explore these relationships, we carried out a laboratory experiment on intact peat soil cores under controlled temperature and water table conditions to determine the impact and interaction of each of these climatic factors on net DOC production. We found a significant interaction (P < 0.001) between temperature, water table draw‐down and net DOC production across the whole soil core (0 to −55 cm depth). This corresponded to an increase in the Q₁₀ (i.e. rise in the rate of net DOC production over a 10 °C range) from 1.84 under high water tables and anaerobic conditions to 3.53 under water table draw‐down and aerobic conditions between −10 and − 40 cm depth. However, increases in net DOC production were only seen after water tables recovered to the surface as secondary changes in soil water chemistry driven by sulphur redox reactions decreased DOC solubility, and therefore DOC concentrations, during periods of water table draw‐down. Furthermore, net microbial consumption of DOC was also apparent at − 1 cm depth and was an additional cause of declining DOC concentrations during dry periods. Therefore, although increased temperature and decreased rainfall could have a significant effect on net DOC release from peatlands, these climatic effects could be masked by other factors controlling the biological consumption of DOC in addition to soil water chemistry and DOC solubility. These findings highlight both the sensitivity of DOC release from ombrotrophic peat to episodic changes in water table draw‐down, and the need to disentangle complex and interacting controls on DOC dynamics to fully understand the impact of environmental change on this system.     

Light availability regulates the response of algae and heterotrophic bacteria to elevated nutrient levels and warming in a northern boreal peatland

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青藏高原泥炭地水位下降促进土壤碳积累的影响机制

酚类物质作为泥炭地重要的碳分解抑制剂,植被作为泥炭地关键的碳输入来源,它们在土壤碳(可溶性有机碳(DOC)等)周转过程中都发挥着重要作用。然而,目前关于植被群落结构、酚类物质以及DOC含量对水位波动的响应存在较大争议。因此,为明确泥炭地水位下降对植被群落结构、酚类物质以及DOC含量的影响并探明三者间的潜在联系,以若尔盖高原泥炭地作为研究对象,选取红原县日干乔地区3处不同地下水位泥炭地(水位由高到低依次为S1(-1.9 cm)、S2(-10 cm)、S3(-19 cm)样地),调查不同水位条件下植被群落结构特征,并探究酚类物质及土壤碳含量对水位波动的响应。结果表明:(1)从S1到S3样地水位下降促进土壤DOC显著增加(P<0.05),土壤总碳从S1到S2显著增加(P<0.05),而从S2到S3无显著差异;(2)泥炭地水位下降促使禾本科(发草Deschampsia cespitosa)、莎草科(木里薹草Carex muliensis、乌拉草Carex meyeriana)植物大量出现,植被群落高度显著增加(P<0.05)。植被群落地上生物量由153.67 g/m~2增加至...     

Carbon turnover in peatland mesocosms exposed to different water table levels

Changes of water table position influence carbon cycling in peatlands, but effects on the sources and sinks of carbon are difficult to isolate and quantify in field investigations due to seasonal dynamics and covariance of variables. We thus investigated carbon fluxes and dissolved carbon production in peatland mesocosms from two acidic and oligotrophic peatlands under steady state conditions at two different water table positions. Exchange rates and CO₂, CH₄ and DOC production rates were simultaneously determined in the peat from diffusive-advective mass-balances of dissolved CO₂, CH₄ and DOC in the pore water. Incubation experiments were used to quantify potential CO₂, CH₄, and DOC production rates. The carbon turnover in the saturated peat was dominated by the production of DOC (10–15 mmol m^–2 d^–1) with lower rates of DIC (6.1–8.5 mmol m^–2 d^–1) and CH₄ (2.2–4.2 mmol m^–2 d^–1) production. All production rates strongly decreased with depth indicating the importance of fresh plant tissue for dissolved C release. A lower water table decreased area based rates of photosynthesis (24–42%), CH₄ production (factor 2.5–3.5) and emission, increased rates of soil respiration and microbial biomass C, and did not change DOC release. Due to the changes in process rates the C net balance of the mesocosms shifted by 36 mmol m^–2 d^–1. According to our estimates the change in C mineralization contributed most to this change. Anaerobic rates of CO₂ production rates deeper in the peat increased significantly by a factor of 2–3.5 (DOC), 2.9–3.9 (CO₂), and 3–14 (CH₄) when the water table was lowered by 30 cm. This phenomenon might have been caused by easing an inhibiting effect by the accumulation of CO₂ and CH₄ when the water table was at the moss surface.     

Macroinvertebrate Responses to Algal and Bacterial Manipulations in Streams

Our study was designed to assess the relative importance of algae and bacteria as sources of energy for stream macroinvertebrates. In one experiment, we manipulated algae by artificially shading six sections in each of two streams, one stream with an open canopy (clear-cut drainage basin) and the other with a closed canopy (forested drainage basin); both streams were in Hubbard Brook Experimental Forest, New Hampshire, USA. Chlorophyll a concentrations were reduced from 0.2 to 0.05 μg/cm² in artificially shaded sections of both streams. However, macroinvertebrates showed no response to these algal manipulations in either the clear-cut or forested stream. Nutrient concentrations (N and P) were low and limiting to primary production in both the clear-cut and forested streams. Additionally, both streams had relatively low macroinvertebrate densities suggesting bottom-up controls were important in macroinvertebrate abundance. However, the forested stream did have higher macroinvertebrate densities presumably because of higher inputs of coarse particulate organic matter from the riparian vegetation. In a second experiment, in Augusta Creek, Michigan, we manipulated both algae and bacteria. To reduce algae, we artificially shaded experimental stream channels so that chlorophyll a was reduced from natural levels of 3.0–5.6 to 0.4–0.7 μg/cm². Half of the shaded channels had dissolved organic carbon (DOC – sucrose) dripped into them to raise DOC levels by 2–3 mg/l and thus stimulate bacterial abundance. Open channels, with higher algal abundance, had higher densities of Ephemerella, but only in November when nymphs were larger. Channels with increased DOC had higher bacterial abundances, higher densities of Chironomidae and lower densities of Heptageniidae. Several other macroinvertebrate taxa that were at relatively low abundance in our samples showed no significant response to these manipulations. Our results suggest that early instar Ephemerella may not rely as heavily on algae as later instars. Also, certain taxa were able to use the heterotrophic microbial community, especially chironomids which increased in numbers when bacterial density increased; thus, the bacterial carbon source may be more important to some stream macroinvertebrates than previous studies have suggested.     

Evidence for older carbon loss with lowered water tables and changing plant functional groups in peatlands

A small imbalance in plant productivity and decomposition accounts for the carbon (C) accumulation capacity of peatlands. As climate changes, the continuity of peatland net C storage relies on rising primary production to offset increasing ecosystem respiration (ER) along with the persistence of older C in waterlogged peat. A lowering in the water table position in peatlands often increases decomposition rates, but concurrent plant community shifts can interactively alter ER and plant productivity responses. The combined effects of water table variation and plant communities on older peat C loss are unknown. We used a full-factorial 1-m³ mesocosm array with vascular plant functional group manipulations (Unmanipulated Control, Sedge only, and Ericaceous only) and water table depth (natural and lowered) treatments to test the effects of plants and water depth on CO₂ fluxes, decomposition, and older C loss. We used Δ¹⁴C and δ¹³C of ecosystem CO₂ respiration, bulk peat, plants, and porewater dissolved inorganic C to construct mixing models partitioning ER among potential sources. We found that the lowered water table treatments were respiring C fixed before the bomb spike (1955) from deep waterlogged peat. Lowered water table Sedge treatments had the oldest dissolved inorganic ¹⁴C signature and the highest proportional peat contribution to ER. Decomposition assays corroborated sustained high rates of decomposition with lowered water tables down to 40 cm below the peat surface. Heterotrophic respiration exceeded plant respiration at the height of the growing season in lowered water table treatments. Rates of gross primary production were only impacted by vegetation, whereas ER was affected by vegetation and water table depth treatments. The decoupling of respiration and primary production with lowered water tables combined with older C losses suggests that climate and land-use-induced changes in peatland hydrology can increase the vulnerability of peatland C stores.     

Long-term water table manipulations alter peatland gaseous carbon fluxes in Northern Michigan

Northern peatland water table position is tightly coupled to carbon (C) cycling dynamics and is predicted to change from shifts in temperature and precipitation patterns associated with global climate change. However, it is uncertain how long-term water table alterations will alter C dynamics in northern peatlands because most studies have focused on short-term water table manipulations. The goal of our study was to quantify the effect of long-term water table changes (~80 years) on gaseous C fluxes in a peatland in the Upper Peninsula of Michigan. Chamber methods were utilized to measure ecosystem respiration (ER), gross primary production (GPP), net ecosystem exchange (NEE), and methane (CH₄) fluxes in a peatland experiencing levee induced long-term water table drawdown and impoundment in relation to an unaltered site. Inundation raised water table levels by approximately ~10 cm and resulted in a decrease in ER and GPP, but an increase of CH₄ emissions. Conversely, the drained sites, with water table levels ~15 cm lower, resulted in a significant increase in ER and GPP, but a decrease in CH₄ emissions. However, NEE was not significantly different between the water table treatments. In summary, our data indicates that long-term water table drawdown and inundation was still altering peatland gaseous C fluxes, even after 80 years. In addition, many of the patterns we found were of similar magnitude to those measured in short-term studies, which indicates that short-term studies might be useful for predicting the direction and magnitude of future C changes in peatlands.     

Short-term productivity responses of algae and bacteria to zooplankton grazing in two freshwater lakes

SUMMARY. 1. The specific productivities of algae and bacteria were measured in short-term (4 day) experiments consisting of enclosures with natural or reduced zooplankton biomass. Experiments were repeated five times over a season in each of two lakes that differed in the background concentration of dissolved organic carbon (DOC).
2. Algal biomass as estimated by chlorophyll a was suppressed in enclosures with ambient grazer levels in six of ten experiments and enhanced in one experiment. Distribution of chlorophyll among net and nanoplankton was not significantly affected by grazing.
3. Relative to enclosures with reduced zooplankton, normal grazer biomass (97–466μg 1⁻¹ dry weight) enhanced specific algal productivity in only one of five experiments in the low DOC take and had no effect in all five experiments in the high DOC lake. The main effects of grazers on algae was through removal of biomass rather than through indirect changes in turnover rate.
4. Between experiments, bacterial density was either unaffected, or mildly enhanced (4–87%) in enclosures with ambient macrozooplankton compared to those with reduced levels. Bacterial productivity and turnover estimated by incorporation of [³H]thymidine into DNA showed different responses across experiments; increasing, declining or remaining the same with grazer minipulation. This variability was not related to differences in dissolved primary production or to background DOC between lakes or experiments. Comparison of bacterial productivities based on thymidine incorporation rates with changes in cell densities indicated that control of bacterial loss processes by macrozooplankton is more important than control of growth rates.     

Processes controlling DOC in pore water during simulated drought cycles in six different UK peats

The effect of episodic drought on dissolved organic carbon (DOC) dynamics in peatlands has been the subject of considerable debate, as decomposition and DOC production is thought to increase under aerobic conditions, yet decreased DOC concentrations have been observed during drought periods. Decreased DOC solubility due to drought-induced acidification driven by sulphur (S) redox reactions has been proposed as a causal mechanism; however evidence is based on a limited number of studies carried out at a few sites. To test this hypothesis on a range of different peats, we carried out controlled drought simulation experiments on peat cores collected from six sites across Great Britain. Our data show a concurrent increase in sulphate (SO₄) and a decrease in DOC across all sites during simulated water table draw-down, although the magnitude of the relationship between SO₄ and DOC differed between sites. Instead, we found a consistent relationship across all sites between DOC decrease and acidification measured by the pore water acid neutralising capacity (ANC). ANC provided a more consistent measure of drought-induced acidification than SO₄ alone because it accounts for differences in base cation and acid anions concentrations between sites. Rewetting resulted in rapid DOC increases without a concurrent increase in soil respiration, suggesting DOC changes were primarily controlled by soil acidity not soil biota. These results highlight the need for an integrated analysis of hydrologically driven chemical and biological processes in peatlands to improve our understanding and ability to predict the interaction between atmospheric pollution and changing climatic conditions from plot to regional and global scales.     

Experimental whole‐lake increase of dissolved organic carbon concentration produces unexpected increase in crustacean zooplankton density

The observed pattern of lake browning, or increased terrestrial dissolved organic carbon (DOC) concentration, across the northern hemisphere has amplified the importance of understanding how consumer productivity varies with DOC concentration. Results from comparative studies suggest these increased DOC concentrations may reduce crustacean zooplankton productivity due to reductions in resource quality and volume of suitable habitat. Although these spatial comparisons provide an expectation for the response of zooplankton productivity as DOC concentration increases, we still have an incomplete understanding of how zooplankton respond to temporal increases in DOC concentration within a single system. As such, we used a whole‐lake manipulation, in which DOC concentration was increased from 8 to 11 mg L^?1 in one basin of a manipulated lake, to test the hypothesis that crustacean zooplankton production should subsequently decrease. In contrast to the spatially derived expectation of sharp DOC‐mediated decline, we observed a small increase in zooplankton densities in response to our experimental increase in DOC concentration of the treatment basin. This was due to significant increases in gross primary production and resource quality (lower seston carbon‐to‐phosphorus ratio; C:P). These results demonstrate that temporal changes in lake characteristics due to increased DOC may impact zooplankton in ways that differ from those observed in spatial surveys. We also identified significant interannual variability across our study region, which highlights potential difficulty in detecting temporal responses of organism abundances to gradual environmental change (e.g., browning).     

泥炭沼泽湿地关键元素地球化学特征及其对碳排放的影响机制

硫、铁是泥炭沼泽湿地（泥炭地）中重要的生源要素,其参与下的生物地球化学过程对泥炭地碳循环意义重大。选取德国中部两处典型的雨养型泥炭地高海拔样点（TBP）和低海拔样点（TSP）,通过原位采集泥炭剖面孔隙水和可溶性气体等,研究了硫、铁元素等地球化学变化规律,结合DOC、甲烷（CH₄）和二氧化碳（CO₂）浓度分布,探讨其对泥炭地碳排放的影响。研究结果表明：（1） TBP中总还原无机硫（TRIS）浓度随深度先增后减,且上部0-87 cm平均浓度远高于87 cm深度以下,上部硫酸盐还原作用强烈。结合上部亚铁、硫化氢（H₂S）浓度分布,得知该范围内H₂S主要是通过微生物硫酸盐还原作用（BSR）生成,同时H₂S在孔隙水扩散过程中易与亚铁结合为硫化亚铁,进而生成稳定的黄铁矿,这一反应过程在约60 cm处减缓。（2） TBP、TSP两处采样点中DOC与亚铁、硫酸盐均有较强相关性,是由于地下水位的波动影响氧化还原程度以及微生物活性。两处采样点DOC均与亚铁呈显著正相关关系,表明铁氧化物在厌氧环境中被还原溶解产生亚铁,与其结合的有机碳被释放到溶液中从而导致DOC浓度的升高。TBP中DOC与硫酸盐呈显著负相关关系,表明硫酸盐作为电子受体被还原的过程中消耗酸度使pH值升高,增强了其中微生物的活性,DOC浓度由此增加。（3） CH₄与硫酸盐、TRIS浓度在剖面上均呈现相反变化趋势,表明硫酸盐输入的增加以及硫酸盐还原活动均会抑制CH₄生成。CO₂/CH₄均大于4,表明硫酸盐作为替代电子受体会使厌氧条件下碳矿化转向多CO₂和少CH₄生成。此外,亚铁对于CH₄生成一定程度上会起到低促高抑的效果,而对于CO₂的生成的影响较弱。表明硫酸盐对于CH₄和CO₂生成的影响高于亚铁。研究着重探究硫、铁等关键元素地下部生物地球化学过程对碳排放的影响机制,研究结果可为泥炭地碳排放核算提供理论支撑。     

Accelerated vegetation succession but no hydrological change in a boreal fen during 20 years of recent climate change

Northern mires (fens and bogs) have significant climate feedbacks and contribute to biodiversity, providing habitats to specialized biota. Many studies have found drying and degradation of bogs in response to climate change, while northern fens have received less attention. Rich fens are particularly important to biodiversity, but subject to global climate change, fen ecosystems may change via direct response of vegetation or indirectly by hydrological changes. With repeated sampling over the past 20 years, we aim to reveal trends in hydrology and vegetation in a pristine boreal fen with gradient from rich to poor fen and bog vegetation. We resampled 203 semi‐permanent plots and compared water‐table depth (WTD), pH, concentrations of mineral elements, and dissolved organic carbon (DOC), plant species occurrences, community structure, and vegetation types between 1998 and 2018. In the study area, the annual mean temperature rose by 1.0°C and precipitation by 46 mm, in 20‐year periods prior to sampling occasions. We found that wet fen vegetation decreased, while bog and poor fen vegetation increased significantly. This reflected a trend of increasing abundance of common, generalist hummock species at the expense of fen specialist species. Changes were the most pronounced in high pH plots, where Sphagnum mosses had significantly increased in plot frequency, cover, and species richness. Changes of water chemistry were mainly insignificant in concentration levels and spatial patterns. Although indications toward drier conditions were found in vegetation, WTD had not consistently increased, instead, our results revealed complex dynamics of WTD as depending on vegetation changes. Overall, we found significant trend in vegetation, conforming to common succession pattern from rich to poor fen and bog vegetation. Our results suggest that responses intrinsic to vegetation, such as increased productivity or altered species interactions, may be more significant than indirect effects via local hydrology to the ecosystem response to climate warming.     

Effects of Experimental Water Table and Temperature Manipulations on Ecosystem CO2 Fluxes in an Alaskan Rich Fen

Peatlands store 30% of the world’s terrestrial soil carbon (C) and those located at northern latitudes are expected to experience rapid climate warming. We monitored growing season carbon dioxide (CO₂) fluxes across a factorial design of in situ water table (control, drought, and flooded plots) and soil warming (control vs. warming via open top chambers) treatments for 2 years in a rich fen located just outside the Bonanza Creek Experimental Forest in interior Alaska. The drought (lowered water table position) treatment was a weak sink or small source of atmospheric CO₂ compared to the moderate atmospheric CO₂ sink at our control. This change in net ecosystem exchange was due to lower gross primary production and light-saturated photosynthesis rather than increased ecosystem respiration. The flooded (raised water table position) treatment was a greater CO₂ sink in 2006 due largely to increased early season gross primary production and higher light-saturated photosynthesis. Although flooding did not have substantial effects on rates of ecosystem respiration, this water table treatment had lower maximum respiration rates and a higher temperature sensitivity of ecosystem respiration than the control plot. Surface soil warming increased both ecosystem respiration and gross primary production by approximately 16% compared to control (ambient temperature) plots, with no net effect on net ecosystem exchange. Results from this rich fen manipulation suggest that fast responses to drought will include reduced ecosystem C storage driven by plant stress, whereas inundation will increase ecosystem C storage by stimulating plant growth.     

Sedge Succession and Peatland Methane Dynamics: A Potential Feedback to Climate Change

Under the warmer climate, predicted for the future, northern peatlands are expected to become drier. This drying will lower the water table and likely result in reduced emissions of methane (CH₄) from these ecosystems. However, the prediction of declining CH₄ fluxes does not consider the potential effects of ecological succession, particularly the invasion of sedges into currently wet sites (open water pools, low lawns). The goal of this study was to characterize the relationship between the presence of sedges in peatlands and CH₄ efflux under natural conditions and under a climate change simulation (drained peatland). Methane fluxes, gross ecosystem production, and dissolved pore water CH₄ concentrations were measured and a vegetation survey was conducted in a natural and drained peatland near St. Charles-de-Bellechasse, Quebec, Canada, in the summer of 2003. Each peatland also had plots where the sedges had been removed by clipping. Sedges were larger, more dominant, and more productive at the drained peatland site. The natural peatland had higher CH₄ fluxes than the drained peatland, indicating that drainage was a significant control on CH₄ flux. Methane flux was higher from plots with sedges than from plots where sedges had been removed at the natural peatland site, whereas the opposite case was observed at the drained peatland site. These results suggest that CH₄ flux was enhanced by sedges at the natural peatland site and attenuated by sedges at the drained peatland site. However, the attenuation of CH₄ flux due to sedges at the drained site was reduced in wetter periods. This finding suggests that CH₄ flux could be decreased in the event of climate warming due to the greater depth to the water table, and that sedges colonizing these areas could further attenuate CH₄ fluxes during dry periods. However, during wet periods, the sedges may cause CH₄ fluxes to be higher than is currently predicted for climate change scenarios.     

ALGAE AS COMPETITORS FOR GLUCOSE WITH HETEROTROPHIC BACTERIA1

Dissolved organic carbon (DOC) constitutes the bulk of organic carbon in aquatic environments. The importance of DOC utilization by mixotrophic algae is unclear since heterotrophic bacteria are regarded as more efficient users. We tested the hypothesis that algae decrease the DOC concentration in the light to lower levels than in darkness resulting in competitive exclusion of heterotrophic bacteria according to the mechanistic competition theory. We investigated (a) the uptake kinetics of glucose as a model substrate by two cultured algae and mixed bacteria populations, (b) the competition for glucose between algae and bacteria in chemostats, (c) the effect of discontinuous glucose supply in chemostats, and (d) the minimum glucose concentrations achieved in cultures of algae and bacteria. Bacteria showed higher specific‐glucose‐uptake rates than algae. In chemostats, algae became extinct in the dark and coexisted in the light where they decreased bacteria to lower densities. Discontinuous glucose supply promoted the algae compared to continuous substrate addition. Several algae consumed glucose to lower concentrations in the dark than in the light and showed lower or equal residual glucose concentrations than bacteria. Residual concentrations were not related to allometric traits (cell volume) and photosynthetic potential (chl content). Overall, the hypothesis was not supported, and mechanisms of competition for DOC obviously differed from those for particulate prey. However, since some algae showed lower or equal residual glucose concentrations than bacteria, algal dark uptake of DOC may be important in deep layers of many waters.     

MIXOTROPHIC ALGAE CONSTRAIN THE LOSS OF ORGANIC CARBON BY EXUDATION1

Algae of various taxonomic groups are capable of assimilating dissolved organic carbon (DOC) from their environments (mixotrophy). Recently, we reported that, with increasing biomass of mixotrophs, heterotrophic bacteria did not increase. We hypothesized that algal uptake of external DOC may outweigh their release of DOC by exudation (H1). Here, we addressed an alternative hypothesis that algae did not assimilate external DOC but constrained the release of DOC (H2). In chemostat experiments, we cultured the mixotrophic Chlamydomonas acidophila Negoro together with heterotrophic bacteria. As external substrates, we used glucose, which was potentially available for both bacteria and algae, or fructose, which was available only for bacteria. We increased the biomass of algae by the stepwise addition of phosphorus. Bacterial biomass did not increase in experiments using glucose or when fructose was offered, suggesting that mechanisms other than algal mixotrophy (H1) kept concentrations of bacteria low. Measured exudation rates (percent extracellular release, PER) of mixotrophic algae (Cd. acidophila, Chlorella protothecoides W. Krüger) were very low and ranged between 1.0% and 3.5% at low and moderately high phosphorus concentrations. In contrast, an obligately phototrophic alga (Chlamydomonas segnis H. Ettl) showed higher exudation rates, particularly under phosphorus limitation (70%). The results support H2. If mixotrophy is considered as a mechanism to recycle organic exudates from near the cell surface, this would explain why algae retained mixotrophic capabilities although they cannot compete with bacteria for external organic carbon.