Chemoautotrophic Carbon Fixation Rates and Active Bacterial Communities in Intertidal Marine Sediments

Chemoautotrophy has been little studied in typical coastal marine sediments, but may be an important component of carbon recycling as intense anaerobic mineralization processes in these sediments lead to accumulation of high amounts of reduced compounds, such as sulfides and ammonium. We studied chemoautotrophy by measuring dark-fixation of ¹³C-bicarbonate into phospholipid derived fatty acid (PLFA) biomarkers at two coastal sediment sites with contrasting sulfur chemistry in the Eastern Scheldt estuary, the Netherlands. At one site where free sulfide accumulated in the pore water right to the top of the sediment, PLFA labeling was restricted to compounds typically found in sulfur and ammonium oxidizing bacteria. At the other site, with no detectable free sulfide in the pore water, a very different PLFA labeling pattern was found with high amounts of label in branched i- and a-PLFA besides the typical compounds for sulfur and ammonium oxidizing bacteria. This suggests that other types of chemoautotrophic bacteria were also active, most likely Deltaproteobacteria related to sulfate reducers. Maximum rates of chemoautotrophy were detected in first 1 to 2 centimeters of both sediments and chemosynthetic biomass production was high ranging from 3 to 36 mmol C m⁻² d⁻¹. Average dark carbon fixation to sediment oxygen uptake ratios were 0.22±0.07 mol C (mol O₂)⁻¹, which is in the range of the maximum growth yields reported for sulfur oxidizing bacteria indicating highly efficient growth. Chemoautotrophic biomass production was similar to carbon mineralization rates in the top of the free sulfide site, suggesting that chemoautotrophic bacteria could play a crucial role in the microbial food web and labeling in eukaryotic poly-unsaturated PLFA was indeed detectable. Our study shows that dark carbon fixation by chemoautotrophic bacteria is a major process in the carbon cycle of coastal sediments, and should therefore receive more attention in future studies on sediment biogeochemistry and microbial ecology.     

Sulfur oxidizers dominate carbon fixation at a biogeochemical hot spot in the dark ocean

Bacteria and archaea in the dark ocean (>200 m) comprise 0.3–1.3 billion tons of actively cycled marine carbon. Many of these microorganisms have the genetic potential to fix inorganic carbon (autotrophs) or assimilate single-carbon compounds (methylotrophs). We identified the functions of autotrophic and methylotrophic microorganisms in a vent plume at Axial Seamount, where hydrothermal activity provides a biogeochemical hot spot for carbon fixation in the dark ocean. Free-living members of the SUP05/Arctic96BD-19 clade of marine gamma-proteobacterial sulfur oxidizers (GSOs) are distributed throughout the northeastern Pacific Ocean and dominated hydrothermal plume waters at Axial Seamount. Marine GSOs expressed proteins for sulfur oxidation (adenosine phosphosulfate reductase, sox (sulfur oxidizing system), dissimilatory sulfite reductase and ATP sulfurylase), carbon fixation (ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCO)), aerobic respiration (cytochrome c oxidase) and nitrogen regulation (PII). Methylotrophs and iron oxidizers were also active in plume waters and expressed key proteins for methane oxidation and inorganic carbon fixation (particulate methane monooxygenase/methanol dehydrogenase and RuBisCO, respectively). Proteomic data suggest that free-living sulfur oxidizers and methylotrophs are among the dominant primary producers in vent plume waters in the northeastern Pacific Ocean.     

Inorganic carbon promotes photosynthesis,growth, and maximum biomass of phytoplankton in eutrophic water bodies

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Rising CO2 Levels Will Intensify Phytoplankton Blooms in Eutrophic and Hypertrophic Lakes

Harmful algal blooms threaten the water quality of many eutrophic and hypertrophic lakes and cause severe ecological and economic damage worldwide. Dense blooms often deplete the dissolved CO₂ concentration and raise pH. Yet, quantitative prediction of the feedbacks between phytoplankton growth, CO₂ drawdown and the inorganic carbon chemistry of aquatic ecosystems has received surprisingly little attention. Here, we develop a mathematical model to predict dynamic changes in dissolved inorganic carbon (DIC), pH and alkalinity during phytoplankton bloom development. We tested the model in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa at different CO₂ levels. The experiments showed that dense blooms sequestered large amounts of atmospheric CO₂, not only by their own biomass production but also by inducing a high pH and alkalinity that enhanced the capacity for DIC storage in the system. We used the model to explore how phytoplankton blooms of eutrophic waters will respond to rising CO₂ levels. The model predicts that (1) dense phytoplankton blooms in low- and moderately alkaline waters can deplete the dissolved CO₂ concentration to limiting levels and raise the pH over a relatively wide range of atmospheric CO₂ conditions, (2) rising atmospheric CO₂ levels will enhance phytoplankton blooms in low- and moderately alkaline waters with high nutrient loads, and (3) above some threshold, rising atmospheric CO₂ will alleviate phytoplankton blooms from carbon limitation, resulting in less intense CO₂ depletion and a lesser increase in pH. Sensitivity analysis indicated that the model predictions were qualitatively robust. Quantitatively, the predictions were sensitive to variation in lake depth, DIC input and CO₂ gas transfer across the air-water interface, but relatively robust to variation in the carbon uptake mechanisms of phytoplankton. In total, these findings warn that rising CO₂ levels may result in a marked intensification of phytoplankton blooms in eutrophic and hypertrophic waters.     

Inorganic carbon fixation by chemosynthetic ectosymbionts and nutritional transfers to the hydrothermal vent host-shrimp Rimicaris exoculata

The shrimp Rimicaris exoculata dominates several hydrothermal vent ecosystems of the Mid-Atlantic Ridge and is thought to be a primary consumer harbouring a chemoautotrophic bacterial community in its gill chamber. The aim of the present study was to test current hypotheses concerning the epibiont''s chemoautotrophy, and the mutualistic character of this association. In-vivo experiments were carried out in a pressurised aquarium with isotope-labelled inorganic carbon (NaH¹³CO₃ and NaH¹⁴CO₃) in the presence of two different electron donors (Na₂S₂O₃ and Fe²⁺) and with radiolabelled organic compounds (¹⁴C-acetate and ³H-lysine) chosen as potential bacterial substrates and/or metabolic by-products in experiments mimicking transfer of small biomolecules from epibionts to host. The bacterial epibionts were found to assimilate inorganic carbon by chemoautotrophy, but many of them (thick filaments of epsilonproteobacteria) appeared versatile and able to switch between electron donors, including organic compounds (heterotrophic acetate and lysine uptake). At least some of them (thin filamentous gammaproteobacteria) also seem capable of internal energy storage that could supply chemosynthetic metabolism for hours under conditions of electron donor deprivation. As direct nutritional transfer from bacteria to host was detected, the association appears as true mutualism. Import of soluble bacterial products occurs by permeation across the gill chamber integument, rather than via the digestive tract. This first demonstration of such capabilities in a decapod crustacean supports the previously discarded hypothesis of transtegumental absorption of dissolved organic matter or carbon as a common nutritional pathway.     

Carbon flux between an estuary and the ocean: a case for outwelling

The classical outwelling hypothesis states that small coastal embayments (e.g. estuaries, wetlands) export their excess production to inshore marine waters. In line with this notion, the present study tested whether the Swartkops estuary acts as source or sink for carbon. To this end, concentrations of dissolved inorganic carbon (DIC), dissolved organic carbon (DOC) and particulate organic carbon (POC) were determined hourly during the first monthly spring and neap tides over one year in the tidal waters entering and leaving the estuary. Each sampling session spanned a full tidal cycle, yielding a total of 936 concentration estimates. Carbon fluxes were calculated by integrating concentrations with water flow rates derived from a hydrodynamic model calibrated for each sampling datum. Over the year, exports to marine waters markedly exceeded imports to the estuary for all carbon species: on the basis of total spring tidal drainage area, 1083 g m^–2 of DIC, 103 g m^–2 of DOC, and 123 g m^–2 of POC left the estuary annually. Total carbon export from the estuary to the ocean amounted to 4755 tonnes, of which 83% was in the inorganic form (DIC). Thus, the bulk of carbon moving in the water column is inorganic - yet, DIC seems to be measured only rarely in most flux studies of this nature. Salt marshes cover extensive areas in this estuary and produce some carbon, particularly DOC, but productivity of the local Spartina species is low (P:B=1.1). Consequently, the bulk of carbon exported from the estuary appears to originate from the highly productive macroinvertebrate and the phytoplankton component and not from the salt marsh plants.     

Constraining carbon sources and growth rates of freshwater microbialites in Pavilion Lake using 14C analysis

This study determined the natural abundance isotopic compositions (¹³C, ¹⁴C) of the primary carbon pools and microbial communities associated with modern freshwater microbialites located in Pavilion Lake, British Columbia, Canada. The Δ¹⁴C of dissolved inorganic carbon (DIC) was constant throughout the water column and consistent with a primarily atmospheric source. Observed depletions in DIC ¹⁴C values compared with atmospheric CO₂ indicated effects due either to DIC residence time and/or inputs of ¹⁴C‐depleted groundwater. Mass balance comparisons of local and regional groundwater indicate that groundwater DIC could contribute a maximum of 9–13% of the DIC. ¹⁴C analysis of microbial phospholipid fatty acids from microbialite communities had Δ¹⁴C values comparable with lake water DIC, demonstrating that lake water DIC was their primary carbon source. Microbialite carbonate was also primarily derived from DIC. However, some depletion in microbialite carbonate ¹⁴C relative to lake water DIC occurred, due either to residence time or mixing with a ¹⁴C‐depleted carbon source. A detrital branch covered with microbialite growth was used to estimate a microbialite growth rate of 0.05 mm year^?1 for the past 1000 years, faster than previous estimates for this system. These results demonstrate that the microbialites are actively growing and that the primary carbon source for both microbial communities and recent carbonate is DIC originating from the atmosphere. While these data cannot conclusively differentiate between abiotic and biotic formation mechanisms, the evidence for minor inputs of groundwater‐derived DIC is consistent with the previously hypothesized biological origin of the Pavilion Lake microbialites.     

Novel Chemoautotrophic Endosymbiosis between a Member of the Epsilonproteobacteria and the Hydrothermal-Vent Gastropod Alviniconcha aff. hessleri (Gastropoda: Provannidae) from the Indian Ocean

The hydrothermal-vent gastropod Alviniconcha aff. hessleri from the Kairei hydrothermal field on the Central Indian Ridge houses bacterium-like cells internally in its greatly enlarged gill. A single 16S rRNA gene sequence was obtained from the DNA extract of the gill, and phylogenetic analysis placed the source organism within a lineage of the epsilon subdivision of the Proteobacteria. Fluorescence in situ hybridization analysis with an oligonucleotide probe targeting the specific epsilonproteobacterial subgroup showed the bacterium densely colonizing the gill filaments. Carbon isotopic homogeneity among the gastropod tissue parts, regardless of the abundance of the endosymbiont cells, suggests that the carbon isotopic composition of the endosymbiont biomass is approximately the same as that of the gastropod. Compound-specific carbon isotopic analysis revealed that fatty acids from the gastropod tissues are all ¹³C enriched relative to the gastropod biomass and that the monounsaturated C₁₆ fatty acid that originates from the endosymbiont is as ¹³C enriched relative to the gastropod biomass as that of the epsilonproteobacterial cultures grown under chemoautotrophic conditions. This fractionation pattern is most likely due to chemoautotrophy based on the reductive tricarboxylic-acid (rTCA) cycle and subsequent fatty acid biosynthesis from ¹³C-enriched acetyl coenzyme A. Enzymatic characterization revealed evident activity of several key enzymes of the rTCA cycle, as well as the absence of ribulose-1,5-bisphosphate carboxylase/oxygenase activity in the gill tissue. The results from anatomic, molecular phylogenetic, bulk and compound-specific carbon isotopic, and enzymatic analyses all support the inference that a novel nutritional strategy relying on chemoautotrophy in the epsilonproteobacterial endosymbiont is utilized by the hydrothermal-vent gastropod from the Indian Ocean. The discrepancies between the data of the present study and those of previous ones for Alviniconcha gastropods from the Pacific Ocean imply that at least two lineages of chemoautotrophic bacteria, phylogenetically distinct at the subdivision level, occur as the primary endosymbiont in one host animal type.     

Picoplankton photosynthesis and diurnal variations in photosynthesis-irradiance relationship in a eutrophic and meso-oligotrophic lake

The abundance and relative importance of autotrophic picoplankton were investigated in two lakes of different trophic status. In the eutrophic lake, measurements of primary production were performed on water samples in situ and in a light incubator three times during the day whereas for the oligotrophic lake, only one measurement of primary production was performed on water samples in the incubator. Dark-carbon losses of phytoplankton from Lake Loosdrecht were investigated in time series. Cell numbers of autotrophic picoplankton in eutrophic Lake Loosdrecht (3.2 × 10⁴ cells ml^–1) were lower than in meso-oligotrophic Lake Maarsseveen (9.8 and 11.4 × 10⁴ cells ml^–1 at the surface and bottom respectively). In the phytoplankton of both lakes the ratio of picoplankton production increased with decreasing light intensity. In Lake Loosdrecht depth-integrated contribution of picoplankton to total photosynthesis was less than 4%. The P-I-relationship showed diurnal variations in light saturated photosynthesis, while light limited carbon uptake remained constant during the day. Dark carbon losses from short-term labelled phytoplankton during the first 12 hours of the night period accounted for 10–25% of material fixed during the preceeding light period.     

Light and dark uptake and loss of 14C: methodological problems with productivity measurements in oceanic waters

Measurements of the uptake and loss of ⁴C in the light and in the dark in the Tasman and Coral Seas have revealed methodological problems with the estimation of productivity in these waters. Rates of productivity estimated without replication, time series incubations and dark controls frequently overestimated the true rates of autotrophic production. The data showed unexpectedly high rates of both uptake and loss in the dark in oligotrophic waters. In oligotrophic oceanic waters, dark incorporation of ¹⁴C sometimes equalled the uptake of ¹⁴C in the light bottle. Rapid uptake of isotope in the dark controls appeared to be the result of rapid bacterial growth and metabolism. This problem was exacerbated by agitation of the sample before or during the incubation. Tropical samples were particularly susceptible to problems arising from the agitation of the samples. Latitudinal gradients of dark uptake and loss were revealed in these incubations. The loss of label during 8–12 hours in the dark (after 12 hr in the light) was as high as 50% in subtropical waters. The loss was frequently unmeasurable (< 10%) in temperate waters. The time course of ¹⁴C uptake indicated active grazing in the bottles and suggested that most of the nighttime losses of label were due to grazing by microheterotrophs. Respiratory losses appeared to be small. Calculated values of the assimilation number (or photosynthetic capacity) which did not correct for dark ¹⁴C uptake were too high to be biochemically realistic. The errors were due to the heterotrophic uptake of label and the lack of dark controls. Rapid release of ¹⁴C in the dark after incubation in the light meant that the estimate of productivity was dependant on the trophic state of the sample and on the period of incubation.     

AMMONIUM STIMULATION OF DARK CARBON FIXATION AS AN INDICATOR OF NITROGEN DEFICIENCY IN PHYTOPLANKTON: POTENTIAL ERRORS CAUSED BY AMMONIUM-OXIDIZING BACTERIA1

Stimulation of dark fixation of carbon by NH₄⁺ is often used as an indicator of phytoplankton N deficiency. This assay is based on the influence of available NH₄⁺ on anaplerotic CO₂ fixation by algae. However, carbon fixation by chemoautotrophic NH₄⁺-oxidizing bacteria may also be stimulated by NH₄⁺ enrichment, a process that can mask the algal response in natural communities. NH₄⁺ addition enhanced dark carbon fixation up to 300%, relative to unamended controls, in organisms collected on a 0.7-μm retention filter in oligotrophic Flathead Lake, Montana, but the effect was not detectable in the presence of nitrapyrin, an inhibitor of NH₄⁺-oxidizing bacteria. Dark carbon fixation was enhanced with addition of NH₄⁺ in organisms retained on 2-μm filters (which should allow passage of most bacteria). NH₄⁺ stimulated dark carbon fixation in N-deficient axenic cultures of Chlamydomonas reinhardtii Dang but not in N-replete cultures in both the presence and absence of nitrapyrin. Application of nitrapyrin or size fractionation treatments, to separate the processes of dark carbon fixation by nitrifiers and phytoplankton, may improve the efficacy of assays using NH₄⁺ stimulation of dark carbon fixation to specifically indicate N deficiency in natural algal communities.     

Ubiquitous Dissolved Inorganic Carbon Assimilation by Marine Bacteria in the Pacific Northwest Coastal Ocean as Determined by Stable Isotope Probing

In order to identify bacteria that assimilate dissolved inorganic carbon (DIC) in the northeast Pacific Ocean, stable isotope probing (SIP) experiments were conducted on water collected from 3 different sites off the Oregon and Washington coasts in May 2010, and one site off the Oregon Coast in September 2008 and March 2009. Samples were incubated in the dark with 2 mM ¹³C-NaHCO₃, doubling the average concentration of DIC typically found in the ocean. Our results revealed a surprising diversity of marine bacteria actively assimilating DIC in the dark within the Pacific Northwest coastal waters, indicating that DIC fixation is relevant for the metabolism of different marine bacterial lineages, including putatively heterotrophic taxa. Furthermore, dark DIC-assimilating assemblages were widespread among diverse bacterial classes. Alphaproteobacteria, Gammaproteobacteria, and Bacteroidetes dominated the active DIC-assimilating communities across the samples. Actinobacteria, Betaproteobacteria, Deltaproteobacteria, Planctomycetes, and Verrucomicrobia were also implicated in DIC assimilation. Alteromonadales and Oceanospirillales contributed significantly to the DIC-assimilating Gammaproteobacteria within May 2010 clone libraries. 16S rRNA gene sequences related to the sulfur-oxidizing symbionts Arctic96BD-19 were observed in all active DIC assimilating clone libraries. Among the Alphaproteobacteria, clones related to the ubiquitous SAR11 clade were found actively assimilating DIC in all samples. Although not a dominant contributor to our active clone libraries, Betaproteobacteria, when identified, were predominantly comprised of Burkholderia. DIC-assimilating bacteria among Deltaproteobacteria included members of the SAR324 cluster. Our research suggests that DIC assimilation is ubiquitous among many bacterial groups in the coastal waters of the Pacific Northwest marine environment and may represent a significant metabolic process.     

Responses of Two Coastal Algae (Skeletonema costatum and Chlorella vulgaris) to Changes in Light and Iron Levels1

Iron (Fe) is essential for phytoplankton growth and photosynthesis, and is proposed to be an important factor regulating algal blooms under replete major nutrients in coastal environments. Here, Skeletonema costatum, a typical red-tide diatom species, and Chlorella vulgaris, a widely distributed Chlorella, were chosen to examine carbon fixation and Fe uptake by coastal algae under dark and light conditions with different Fe levels. The cellular carbon fixation and intracellular Fe uptake were measured via ¹⁴C and ⁵⁵Fe tracer assay, respectively. Cell growth, cell size, and chlorophyll-α concentration were measured to investigate the algal physiological variation in different treatments. Our results showed that cellular Fe uptake proceeds under dark and the uptake rates were comparable to or even higher than those in the light for both algal species. Fe requirements per unit carbon fixation were also higher in the dark resulting in higher Fe: C ratios. During the experimental period, high Fe addition significantly enhanced cellular carbon fixation and Fe uptake. Compared to C. vulgaris, S. costatum was the common dominant bloom species because of its lower Fe demand but higher Fe uptake rate. This study provides some of the first measurements of Fe quotas in coastal phytoplankton cells, and implies that light and Fe concentrations may influence the phytoplankton community succession when blooms occur in coastal ecosystems.     

On the distribution and importance of picocyanobacteria in a boreal inshore area (Kiel Bight, Western Baltic)

From April to October 1986 abundance and vertical distributionof picocyanobacteria were studied at four stations in Kiel Fjordand Kiel Bight. Both picocyanobacteria and autotrophic, eukaryoticpicoplankton cell numbers were estimated by epifluorescencemicroscopy whereas larger phytoplankton (>3 µm) wasenumerated by the Utermöhi settling technique. Picocyanobactenacell numbers peaked in July and August near the water surface(1.4–2.6 x 10⁸ cells l^–1). Although picocyanobacteriaabundance increased from the outer Kiel Bight to the more eutrophicinner stations of Kiel Fjord, their contribution to total phytoplanktonbiomass decreased. During summer up to 52% of phytoplanktoncarbon and up to 97% of autotrophic picoplankton carbon werecontributed by picocyanobacteria. Therefore picocyanobacteriaare an important component of the summer phytoplankton communityin boreal inshore waters, too.     

Depleted methane-derived carbon in waters of Lake Baikal,Siberia

Results of hydrochemical and stable isotope measurements during the ice-breaking period on Lake Baikal indicate an apparent lack of relationship between measured δ¹³C of dissolved inorganic carbon (DIC) and phytoplankton below the trophogenic layer. While planktonic values of −31.7 to −33.5‰ are within a typical lacustrine range, the δ¹³C values of DIC turned out to be very negative, from −28.9 to −35.6‰. These isotopic values of DIC appear to be associated with oxidation of methane that accumulated during winter ice cover period. At the time of sampling, however, the observed depletion did not affect the phytoplankton/DIC fractionation relationship, because the difference between phytoplankton and DIC (−20 to −22‰ in surface waters) lies within the expected range of the fractionation coefficient. By analogy with small lakes, we explain this lack of relationship by the time lag between peak productivity and peak methane oxidation. Our interpretation of the Baikal DIC isotopic signature is consistent with methanogenesis in bottom sediments and with the known presence of widespread unstable gas hydrates and active methane seeps on the lake floor. Our findings suggest that methane is an important component of the Baikal carbon cycle, that late winter concentrations of methane in Baikal under ice may be 3–4 orders of magnitude higher than previously reported values for summer, and that the lake may be emitting a significant amount of methane to the atmosphere.     

Particulate organic and dissolved inorganic carbon stable isotopic compositions in Taylor Valley lakes,Antarctica: the effect of legacy

In perennially ice-covered lakes of Taylor Valley, Antarctica, “legacy”, a carryover of past ecosystem events, has primarily been discussed in terms of nutrient and salinity concentrations and its effect on the current ecology of the lakes. In this study, we determine how residual pools of ancient carbon affect the modern carbon abundance and character in the water columns of Lakes Fryxell, Hoare, and Bonney. We measure the stable carbon isotopic compositions and concentrations of particulate organic carbon (POC) and dissolved inorganic carbon (DIC) in the water column of these lakes over four seasons (1999–2002). These data are presented and compared with all the previously published Taylor Valley lacustrine carbon stable isotopic data. Our results show that the carbon concentrations and isotopic compositions of the upper water columns of those lakes are controlled by modern processes, while the lower water columns are controlled to varying degrees by inherited carbon pools. The water column of the west lobe of Lake Bonney is dominated by exceptionally high concentrations of DIC (55,000–75,000 μmol l⁻¹) reflecting the long period of ice-cover on this lake. The east lobe of Lake Bonney has highly enriched δ¹³C_DIC values resulting from paleo-brine evaporation effects in its bottom waters, while its high DIC concentrations provide geochemical evidence that its middle depth waters are derived from West Lake Bonney during a hydrologically connected past. Although ancient carbon is present in both Lake Hoare and Lake Fryxell, the δ¹³C_DIC values in bottom waters suggest dominance by modern primary productivity-related processes. Anaerobic methanogenesis and methanotrophy are also taking place in the lower water column of Lake Fryxell with enough methane, oxidized anaerobically, to contribute to the DIC pool. We also show how stream proximity and high flood years are only a minor influence on the carbon isotopic values of both POC and DIC. The Taylor Valley lake system is remarkably stable in both inter-lake and intra-lake carbon dynamics. Handling editor: K. Martens     

Production of new organic carbon and its distribution between autotrophic picoplankton, bacteria, extracellular organic carbon and phytoplankton in an upland lake

• 1 Picoplankton community production (0.2–2μm) was investigated over 3 months, June-September 1991, in Llyn Padarn, a mesotrophic upland lake in north Wales.
• 2 The picoplankton was differentiated into autotrophic algae (<1–3μm) and heterotrophic bacteria (<0.2–1 μm) using differential filtration through a 1 μm pore size Nuclepore filter.
• 3 Efficient separation of these distinct metabolic constituents of picoplankton was obtained. A good correlation (r= 0.81, P < 0.001) was found between physical separation of bacterial and picoalgal cells from fluorescence microscopy and the distribution of heterotrophic metabolic activity between different cell size fractions measured by uptake of ¹⁴C-glucose.
• 4 Picoplankton community production was differentiated into the ‘absolute’ autotrophic production by picoalgae, corrected for overestimation due to retention of bacteria with the picoalgae, and the heterotrophic component, bacterial uptake of ‘extracellular organic carbon’ (EOC), derived from the entire phytoplankton community.
• 5 The heterotrophic contribution to picoplankton community production ranged from 88 to 1%, mean value 55% of total. Autotrophic picoplankton production was dominant in June and July, but in August and September heterotrophic uptake of EOC was the major input to picoplankton community production.
• 6 During the 3 months, the mean contributions to plankton production were autotrophic picoplankton 10.3%, heterotrophic bacterial uptake of EOC 9.7%, EOC in lake water 11.6% and phytoplankton (>3μm) 68.3%.
• 7 Bacteria accounted for about half the picopfankton community production via uptake of EOC. Thus although autotrophic picoplankton were ubiquitous, it is likely that their contribution via primary production to the carbon balance of planktonic environments has been overestimated in previous studies.

     

INFLUENCE OF UV-B RADIATION ON NITROGEN UTILIZATION BY A NATURAL ASSEMBLAGE OF PHYTOPLANKTON

A 7‐day mesocosm experiment was conducted in July 1996 to investigate the effects of ambient UV‐B radiation (UVBR) exclusion and two UVBR enhancements above ambient levels on NO₃^?, NH₄⁺ and urea utilization in a natural plankton community (<240 μm) from the Lower St. Lawrence Estuary. The phytoplankton community was dominated by diatoms during the first 3 days and, afterward, by flagellates and dinoflagellates. The results of 4‐h incubations just below the water surface show that, compared with ambient UVBR conditions, UVBR exclusion generally increased NO₃^?, NH₄⁺, and urea uptakes. During the last 4 days of the experiment, the percent increase in the specific uptake rate of urea under excluded UVBR conditions varied between 17% and 130% and was a linear function of the ambient UVBR dose removed. During the first 3 days, the phytoplankton community dominated by diatoms was able to withstand UVBR enhancements without any perceptible effect on nitrogen uptake. However, during the post‐diatom bloom period, UVBR enhancements resulted in decreases in NO₃^?, NH₄⁺, and urea uptake compared with ambient UVBR conditions. The reduction of urea uptake under UVBR enhancements during the last 3 days varied between 23% and 64% and was linearly related to the enhanced UVBR dose. However, the different UVBR treatments did not affect the internal organic nitrogen composition (internal urea, free amino acids, and proteins) of the phytoplankton community experiencing vertical mixing in the mesocosms. The discrepancy between short‐term uptake measurements at the surface and long‐term effects in the mesocosms emphasizes the importance of vertical mixing on UVBR effects in natural ecosystems. This suggests that an increase in ambient UVBR would have a minimal effect on nitrogen utilization by natural phytoplankton assemblages if these are vertically mixed.     

Some Observations on the Irradiance and Carbon Fixation in Grane Langsø

Photosynthetically active radiation (PAR) was registered daily in 0.25 m and 11.25 m in 3 years. Upwards irradiance of green light was 1 % of downward irradiance. Incubator ¹⁴C experiments showed that the phytoplankton enhanced its carbon uptake substantially with increasing concentrations of the carrier CO₂ in the ampoules. Severe carbon limitation of photosynthesis occurred in spring and summer 1961, where the real carbon fixation was only about 34 % of that calculated by the usual procedure. Utilization of light and DIC by the phytoplankton and its compensation depth was determined. Photoinhibition occurred down to 10–11 m, and net primary production was nearly always positive in any depth down to the lake bottom.