Use of phytoplankton-derived dissolved organic carbon by different types of bacterioplankton

Phytoplankton and heterotrophic prokaryotes are major components of the microbial food web and interact continuously: heterotrophic prokaryotes utilize the dissolved organic carbon derived from phytoplankton exudation or cell lysis (DOCp), and mineralization by heterotrophic prokaryotes provides inorganic nutrients for phytoplankton. For this reason, these communities are expected to be closely linked, although the study of the interactions between them is still a major challenge. Recent studies have presented interactions between phytoplankton and heterotrophic prokaryotes based on coexistence or covariation throughout a time-series. However, a real quantification of the carbon flow within these networks (defined as the interaction strength, IS) has not been achieved yet. This is critical to understand the selectivity degree of bacteria responding to specific algal DOCp. Here we used microautoradiography to quantify the preferences of the major heterotrophic prokaryote phylogenetic groups on DOC derived from several representative phytoplankton species, and expressed these preferences as an IS value. The distribution of the ISs was not random but rather skewed towards weak interactions, in a similar way as the distributions described for stable complex non-microbial ecosystems, indicating that there are some cases of high specificity on the use of specific algal DOCp by some bacterial groups, but weak interactions are more common and may be relevant as well. The variety of IS patterns observed supports the view that the vast range of different resources (different types of organic molecules) available in the sea selects and maintains the high levels of diversity described for marine bacterioplankton.     

Temperature regulation of marine heterotrophic prokaryotes increases latitudinally as a breach between bottom‐up and top‐down controls

Planktonic heterotrophic prokaryotes make up the largest living biomass and process most organic matter in the ocean. Determining when and where the biomass and activity of heterotrophic prokaryotes are controlled by resource availability (bottom‐up), predation and viral lysis (top‐down) or temperature will help in future carbon cycling predictions. We conducted an extensive survey across subtropical and tropical waters of the Atlantic, Indian and Pacific Oceans during the Malaspina 2010 Global Circumnavigation Expedition and assessed indices for these three types of controls at 109 stations (mostly from the surface to 4,000 m depth). Temperature control was approached by the apparent activation energy in eV (ranging from 0.46 to 3.41), bottom‐up control by the slope of the log‐log relationship between biomass and production rate (ranging from ?0.12 to 1.09) and top‐down control by an index that considers the relative abundances of heterotrophic nanoflagellates and viruses (ranging from 0.82 to 4.83). We conclude that temperature becomes dominant (i.e. activation energy >1.5 eV) within a narrow window of intermediate values of bottom‐up (0.3–0.6) and top‐down 0.8–1.2) controls. A pervasive latitudinal pattern of decreasing temperature regulation towards the Equator, regardless of the oceanic basin, suggests that the impact of global warming on marine microbes and their biogeochemical function will be more intense at higher latitudes. Our analysis predicts that 1°C ocean warming will result in increased biomass of heterotrophic prokaryoplankton only in waters with <26°C of mean annual surface temperature.     

Dissolved Oxygen Saturation Controls PAH Biodegradation in Freshwater Estuary Sediments

Polycyclic aromatic hydrocarbons (PAHs) are common contaminants in terrestrial and aquatic environments and can represent a significant constituent of the carbon pool in coastal sediments. We report here the results of an 18-month seasonal study of PAH biodegradation and heterotrophic bacterial production and their controlling biogeochemical factors from 186 sediment samples taken in a tidally influenced freshwater estuary. For each sampling event, measurements were averaged from 25–45 stations covering ∼250 km². There was a clear relationship between bacterial production and ambient temperature, but none between production and bottom water dissolved oxygen (DO) % saturation or PAH concentrations. In contrast with other studies, we found no effect of temperature on the biodegradation of naphthalene, phenanthrene, or fluoranthene. PAH mineralization correlated with bottom water DO saturation above 70% (r² > 0.99). These results suggest that the proportional utilization of PAH carbon to natural organic carbon is as much as three orders of magnitude higher during cooler months, when water temperatures are lower and DO % saturation is higher. Infusion of cooler, well-oxygenated water to the water column overlying contaminated sediments during the summer months may stimulate PAH metabolism preferentially over non-PAH organic matter.     

Contribution of Sediment Respiration to Summer CO2 Emission from Low Productive Boreal and Subarctic Lakes

We measured sediment production of carbon dioxide (CO₂) and methane (CH₄) and the net flux of CO₂ across the surfaces of 15 boreal and subarctic lakes of different humic contents. Sediment respiration measurements were made in situ under ambient light conditions. The flux of CO₂ between sediment and water varied between an uptake of 53 and an efflux of 182 mg C m⁻² day⁻¹ from the sediments. The mean respiration rate for sediments in contact with the upper mixed layer (SedR) was positively correlated to dissolved organic carbon (DOC) concentration in the water (r² = 0.61). The net flux of CO₂ across the lake surface [net ecosystem exchange (NEE)] was also closely correlated to DOC concentration in the upper mixed layer (r² = 0.73). The respiration in the water column was generally 10-fold higher per unit lake area compared to sediment respiration. Lakes with DOC concentrations <5.6 mg L⁻¹ had net consumption of CO₂ in the sediments, which we ascribe to benthic primary production. Only lakes with very low DOC concentrations were net autotrophic (<2.6 mg L⁻¹) due to the dominance of dissolved allochthonous organic carbon in the water as an energy source for aquatic organisms. In addition to previous findings of allochthonous organic matter as an important driver of heterotrophic metabolism in the water column of lakes, this study suggests that sediment metabolism is also highly dependent on allochthonous carbon sources.     

Microbial loop malfunctioning in the annual sea ice at Terra Nova Bay (Antarctica)

We investigated organic carbon quantity and biochemical composition, prokaryotic abundance, biomass and carbon production in the annual and platelet sea ice of Terra Nova Bay (Antarctica), as well as the downward fluxes of organic matter released by melting ice during early spring. Huge amounts of biopolymeric C accumulated in the bottom layer of the ice column concomitantly with the early spring increase in sympagic algal biomass. Such organic material, mostly accounted for by autotrophic biomass, was characterised by a high food quality and was rapidly exported to the sea bottom during sea ice melting. Prokaryote abundance (up to 1.3 × 10⁹ cells L⁻¹) and extracellular enzymatic activities (up to 24.3 μM h⁻¹ for amino-peptidase activity) were extremely high, indicating high rates of organic C degradation in the bottom sea ice. Despite this, prokaryote C production values were very low (range 5–30 ng C L⁻¹ h⁻¹), suggesting that most of the degraded organic C was not channelled into prokaryote biomass. In the platelet ice, we found similar organic C concentrations, prokaryote abundance and biomass values and even higher extracellular enzymatic activities, but values of prokaryote C production (range 800–4,200 ng C L⁻¹ h⁻¹) were up to three orders of magnitude higher than in the intact bottom sea ice. Additional field and laboratory experiments revealed that the dissolved organic material derived from algae accumulating in the bottom sea ice significantly reduced prokaryote C production, suggesting the presence of a potential allopathic control of sympagic algae on prokaryote growth. This article belongs to a special topic: Five articles on Sea-ice communities in Terra Nova Bay (Ross Sea), coordinated by L. Guglielmo and V. Saggiomo, appear in this issue of Polar Biology. The studies were conducted in the frame of the National Program of Research in Antarctica (PNRA) of Italy.     

Ultra- and microplankton assemblages as indicators of trophic status in a Mediterranean lagoon

The seasonal abundance distribution of heterotrophic prokaryotes, pico- and nanophytoplankton, was investigated in connection with environmental variables and microplankton abundance at five stations in Ghar El Melh Lagoon (northeastern Tunisia). Flow cytometry analysis of ultraplankton resolved (i) five heterotrophic prokaryote groups labelled LNA1, LNA2 (low nucleic acid content), HNA1, HNA2 and HNA3 (high nucleic acid content) and (ii) at least 14 ultraphytoplankton groups assigned to picoeukaryotes, picoprokaryotes, nanoeukaryotes, cryptophyte-like cells and some unknown communities. Redundancy analysis (RDA) revealed (i) autumn-summer outbreaks of heterotrophic prokaryotes dominated by HNA groups and (ii) winter-summer proliferation of ultraphytoplankton dominated by nanophytoplankton groups. Generalized additive models (GAM) highlighted the role of (i) water temperature and orthophosphate concentrations in heterotrophic prokaryote distribution and (ii) water temperature and salinity in ultraphytoplankton abundance variation. Based on Spearman's rank correlation, significant negative correlations were established between ultra- and microplankton communities suggesting that, through grazing pressure, microplankton may be behind the drastic decrease in ultraplankton abundances in spring.     

Major Effect of Hydrogen Peroxide on Bacterioplankton Metabolism in the Northeast Atlantic

Reactive oxygen species such as hydrogen peroxide have the potential to alter metabolic rates of marine prokaryotes, ultimately impacting the cycling and bioavailability of nutrients and carbon. We studied the influence of H₂O₂ on prokaryotic heterotrophic production (PHP) and extracellular enzymatic activities (i.e., β-glucosidase [BGase], leucine aminopeptidase [LAPase] and alkaline phosphatase [APase]) in the subtropical Atlantic. With increasing concentrations of H₂O₂ in the range of 100–1000 nM, LAPase, APase and BGase were reduced by up to 11, 23 and 62%, respectively, in the different water layers. Incubation experiments with subsurface waters revealed a strong inhibition of all measured enzymatic activities upon H₂O₂ amendments in the range of 10–500 nM after 24 h. H₂O₂ additions also reduced prokaryotic heterotrophic production by 36–100% compared to the rapid increases in production rates occurring in the unamended controls. Our results indicate that oxidative stress caused by H₂O₂ affects prokaryotic growth and hydrolysis of specific components of the organic matter pool. Thus, we suggest that oxidative stress may have important consequences on marine carbon and energy fluxes.     

Tidal impact on planktonic primary and bacterial production in the German Wadden Sea

 Tidal variation of biological parameters was studied at three anchor stations in selected inlet channels of the northern German Wadden Sea in May and July 1994. Concentrations of bacteria, chlorophyll a and suspended matter as well as primary and bacterial production were assessed over a period of 25 h in the surface and in the bottom water. Diurnal variation in primary production was found both under in situ light conditions and under constant illumination. Tidal turbulence caused the introduction of detritus, bacteria and pigments from the sediment into the water column. The impact of sediment resuspension was most evident in the bottom water, leading to tidally oscillating bacterial production rates which were high during high stream velocity and low during the slack times. Estimations of the areal daily phytoplankton production and corresponding bacterial carbon demands were unbalanced. Primary production accounted for only 25–45% of the total bacterial carbon requirement. This discrepancy is due to the shallow euphotic depth in the Wadden Sea, allowing net primary production only in the upper 2–3 m of the water column, while the relatively high levels of bacterial activity do not show a vertical decline. Assuming that the specific biological activities in the water columns over the tidal flats are similar to those found in the inlet channels, it was found that production processes dominate in shallow areas whereas decomposition processes dominate in the deep channels. Moreover, the predominance of heterotrophic processes in the inlet channels means that additional organic carbon sources must contribute to the heterotrophic metabolism in the deep parts of the Wadden Sea, and that the horizontal flux of material is important in this turbid mesotidal ecosystem. Received: 18 April 1998 / Accepted: 12 November 1998     

Micro-organism activity in aggregate layers in shallow eutrophic brackish water as influenced by wind induced mixing; an experimental approach

In the shallow eutrophic waters of the Darss-Zingst-ecosystem (Baltic Sea) wind induced mixing of the water column, and resuspension of sediment is an important phenomenon. Incubation experiments (7 days) with natural water were performed in the laboratory under simulated mixing conditions in order to assess the effects on abundance, biomass and activity of bacteria, heterotrophic nanoflagellates and ciliates in the water column and upper bottom layers. Under calm mixing conditions a bottom layer developed, rich in organic aggregates. Bacterial biomass increased through increase of bacteria abundance and doubled cell volume. Consequently, biomass of flagellates and, more pronounced, of ciliates increased. The bottom layer, comprising only 15% of the incubated volume, containedca. 60% of the bacterial biomass and ca. 55% of production, 75 and 95% of flagellate biomass, and ca. 70% of ciliate biomass. It is concluded that bottom layers rich in organic aggregates, although not being present permanently, play an important role in the shallow Darss-Zingst-ecosystem.     

Microbial carbon processing in oligotrophic Lake Lucerne (Switzerland): results of in situ <Superscript>13</Superscript>C-labelling studies

Although lakes play a major role in the storage of organic carbon, processes involved are not yet very well characterized, especially for oligotrophic lakes. Whether a lake functions as a net source or sink for carbon depends on relative rates of primary production, inputs of terrestrial organic matter and respiration. The microbial community will affect the efficiency of carbon cycling and thereby potential carbon storage. Because the organic matter fluxes are smaller in oligotrophic lakes they have been studied less intensively with respect to their carbon cycling compared to eutrophic lakes. Whether they play an appreciable role in freshwater carbon cycling relies on unraveling primary and secondary production. Here we present the results from such a study in oligotrophic Lake Lucerne, Switzerland. Based on in situ carbon isotopic labelling experiments using dark, glucose-labelled and transparent, DIC-labelled bottles positioned at different depths in the water column, we conclude that even though the photic zone was very deep, integrated primary productivity was consistently low. The carbon processing efficiency of the heterotrophic producers was such that photosynthesized organic matter was fully consumed, even during times of maximum productivity. This implies that the heterotrophic producers were well adapted to rapidly respond to a temporary increase in primary productivity, which is in line with calculated bacterial growth efficiencies in the surface water layer. Highest glucose-based productivity, as a measure of the heterotrophic potential, was observed in the deepest parts of the water column. Chemoautotrophy was shown at 60 m water depth and is of relatively minor importance for overall fluxes. Mixotrophy was recognized as a strategy to keep up production when light conditions become less favorable for autotrophic growth. A mesocosm experiment earlier in the year indicated lower primary production, which agrees well with the timing of this experiment preceding the annual spring bloom. During the low-productivity season the coupling between phytoplankton and bacterial production was much weaker and potentially more organic matter could escape recycling at that time, although quantitatively fluxes remained very low.     

Pelagic microbial activity in the Northeast Water polynya, summer 1992

 Heterotrophic activity and related measures of pelagic microorganisms in the Northeast Water (NEW) polynya, the largest and northernmost summer polynya in the Arctic, were studied during the Polar Sea cruise of July/August 1992 at stations spanning a range of ice conditions. Utilization (incorporation and respiration) of ¹⁴C-labelled amino acids was measured at in situ temperature as a proxy for microheterotrophic activity in samples from the chlorophyll maxima, intermediate water depths, and the benthic boundary layer. Total activity and bacterial abundance (measured by epifluorescence microscopy) were highest in the sub-zero surface water layer that dominates the northeast Greenland shelf, and particularly in areas most influenced by ice-edge processes and lateral advection. In this cold-water layer activity correlated strongly with particulate organic carbon (POC), to a lesser degree with phytoplankton-derived pigments and bacterial abundance, but not with median bacterial cell size. No significant correlations were detected in the warmer, Atlantic-influenced deep waters of the regional trough system where POC concentrations and other parameters were low. In comparison with temperate environments, pelagic heterotrophic activity in the NEW polynya appears to be low, a finding that bears upon the fate of dissolved organic matter and the efficient supply of particulate food to the benthos in this unique polar setting. Received: 3 December 1995/Accepted:29 May 1996     

Fish Farming Effects on Chemical and Microbial Variables of the Water Column: A Spatio-temporal Study Along the Mediterranean Sea

The effects of fish farming on chemical and biological variables of the water column were investigated in three coastal locations along the Mediterranean (Spain, Italy and Greece). Analyses of nutrients, chlorophyll a, particulate organic carbon and nitrogen (POC and PON), heterotrophic bacteria and cyanobacteria (Synechococcus) were carried out on integrated samples (0–30 m depth) taken at various distances from fish farms and the respective reference sites. At one of these sites, water samples were collected by means of Niskin bottles at 09:00, 13:00 and 16:00 h and at three different depths (0 m, 10 m and bottom). Integrated sampling showed no significant changes with distance in any of the variables measured, whereas all variables except PON showed significant changes between the three locations studied. However, changes were found in samples taken by Niskin bottles, with increased concentrations of NH₄⁺, POC, PON and decreased cyanobacteria densities at the surface layer in the immediate vicinity of the cages. Samples taken at the cages in the afternoon showed increased PO₄³⁻ concentrations in comparison to those taken in the morning. Dilution and grazing are probably both responsible for the lack of detectable signs of eutrophication.     

Mesopelagic microbial carbon production correlates with diversity across different marine particle fractions

The vertical flux of marine snow particles significantly reduces atmospheric carbon dioxide concentration. In the mesopelagic zone, a large proportion of the organic carbon carried by sinking particles dissipates thereby escaping long term sequestration. Particle associated prokaryotes are largely responsible for such organic carbon loss. However, links between this important ecosystem flux and ecological processes such as community development of prokaryotes on different particle fractions (sinking vs. non-sinking) are yet virtually unknown. This prevents accurate predictions of mesopelagic organic carbon loss in response to changing ocean dynamics. Using combined measurements of prokaryotic heterotrophic production rates and species richness in the North Atlantic, we reveal that carbon loss rates and associated microbial richness are drastically different with particle fractions. Our results demonstrate a strong negative correlation between prokaryotic carbon losses and species richness. Such a trend may be related to prokaryotes detaching from fast-sinking particles constantly enriching non-sinking associated communities in the mesopelagic zone. Existing global scale data suggest this negative correlation is a widespread feature of mesopelagic microbes.Subject terms: Microbial ecology, Biogeochemistry     

THE BOTTOM-ICE MICROALGAL COMMUNITY FROM ANNUAL ICE IN THE INSHORE WATERS OF EAST ANTARCTICA1

The structure, productivity and heterotrophic potential of an extensive microalgal community growing on the underside of sea ice near the Australian Antarctic Station of Casey, are described. Underwater observations made near the Australian Antarctic stations of Davis and Mawson are also reported. This community develops during September, is largely suspended from the bottom surface of annual sea ice and often extends into the underlying water column as conspicuous strands up to 15 cm long. The algal community structure in the strands is dominated by an unidentified tube diatom belonging to the Amphipleura/Berkeleya group and chains of a species of Entomoneis cf. Amphiprora paludosa var. hyperborea (Grunow) Cleve. Unlike previously described bottom ice environments, a brash ice layer under the hard sea ice is absent. Living cells, predominantly Nitzschia frigida Grunow, also occur in microbrine channels in the bottom 3 cm of the ice. Maximal primary production rates of 81 μg C · L^-1· h^-1 occurred during November, then began declining near the end of December. Minimal rates (2.8 μg C · L^-1· h^-1) were reached in mid-January and coincided with changes in the physical structure of the sea ice and in the stability of the water column. An abundant epibacterial community associated with the microalgal strands assimilated ³H-labelled amino acids suggesting significant heterotrophic recycling of dissolved organic matter. Turnover times of assimilated amino acids in the bottom ice community averaged 55 h during November while negligible turnover of these substrates occurred in the water column 1.5 m below the ice. These bottom ice communities have higher primary productivity than typical brash ice communities; they are also accessible to marine herbivores and so may be more important to the Antarctic marine food chain than previously supposed.     

Surface microlayers on temperate lowland lakes

At the air–water interface material, organisms accumulate and form a thin layer of organic and inorganic material called the surface microlayer (SML). In order to investigate the development, composition, and metabolism of SML on lakes, samples were collected using a screen sampler along with subsurface water (SSW) in an eutrophic and a mesotrophic lake from April to September 2007. Wind, solar irradiance, and lake temperature were followed continuously. Samples were analyzed for organic and inorganic compounds as well as for photosynthesis and respiration. Most compounds were enriched in the SML relative to the SSW. Enrichment was small, however, probably because sampling was performed on nonslick areas. Most compounds correlated closely between the SML and the SSW, confirming the hypothesis that most SML material originates from the bulk water. Correlations were strongest in the eutrophic lake, probably because external sources had a greater effect on SML concentrations in the mesotrophic lake. Enrichment of compounds and metabolic rates in the SML had similar seasonality and dependency of climatic conditions in the two lakes, suggesting common regulating mechanisms of enrichment and production. Enrichment factors of several compounds were higher at low bulk water concentrations, suggesting that atmospheric deposition then contributed relatively more to concentrations in the SML. Increasing temperature significantly decreased SML enrichment of TOC (total organic carbon), related to changes in TOC composition and higher heterotrophic activity, while wind and solar irradiance had no pronounced enrichment effect on any compound. Net photosynthesis was significantly lower in the SML, experiencing photoinhibition in one-third of the samples. In contrast, respiration was much elevated in the SML. Nonetheless, respiration in the SML never contributed by more than 0.3% of water column respiration, but the combination of enhanced degradation rates of organic carbon in the SML and strong interaction with water below suggests that the SML, nonetheless, may play an important role in degradation of refractory organic carbon. Combining these results, we found that the SML of nonslicked areas on lakes are enriched in organic and inorganic pools and constitute a strong heterotrophic environment, albeit of minor importance for whole lake pelagic metabolism. Handling editor: D. Ryder     

Seasonal Depth-Related Gradients in Virioplankton: Standing Stock and Relationships with Microbial Communities in Lake Pavin (France)

This study presents a depth-related survey of virioplankton abundance in Lake Pavin (Massif Central, France), in relation to the abundances of heterotrophic prokaryotes, picocyanobacteria (Pcy), autotrophic picoeukaryotes (Peu), and of autotrophic (ANF) and heterotrophic (HNF) nanoflagellates. The sampling strategy was designed to be representative of the physico-chemical gradients of the whole water column of the lake, and the seasonal variability as well. In mixolimnic surface waters, all communities were present and viral abundance peaked in summer and autumn. Viral abundance was significantly correlated (p < 0.001) with Pcy, Peu, and ANF, indicating that cyanophages and perhaps other phytoplankton viruses represent a significant pool of viral standing stocks in the mixolimnion of Lake Pavin. Microautotrophs were absent in the deep monimolimnic water masses, where viruses and heterotrophic prokaryotes exhibited highest seasonal abundances in summer and/or autumn and were significantly correlated (p < 0.001) to each other. This indicates that the anoxic monimolimnion of Lake Pavin is an exclusive habitat for viruses and heterotrophic prokaryotes. We conclude that in this habitat, host availability is prevalent over other factors (temperature, oxygen, nutrients, grazers) in favoring viral proliferation.     

Bottom–up carbon subsidies and top–down predation pressure interact to affect aquatic food web structure

Human impacts such as eutrophication, overexploitation and climate change currently threaten future global food and drinking water supplies. Consequently, it is important that we understand how anthropogenic resource (bottom–up) and consumer (top–down) manipulations affect aquatic food web structure and production. Future climate changes are predicted to increase the inputs of terrestrial dissolved organic carbon to lakes. These carbon subsidies can either increase or decrease total basal production in aquatic food webs, depending on bacterial competition with phytoplankton for nutrients. This study examines the effects of carbon subsidies (bottom–up) on a pelagic community exposed to different levels of top–down predation. We conducted a large scale mesocosm experiment in an oligotrophic clear water lake in northern Sweden, using a natural plankton community exposed to three levels of glucose addition (0, 420 and 2100 μg C l^?1 total added glucose) and three levels of young‐of‐the‐year perch Perca fluviatilis density (0, 0.56 and 2 individuals m^?3). Bacterioplankton production doubled with glucose addition, but phytoplankton production was unaffected, in contrast to previous studies that have manipulated carbon, nutrients or light simultaneously. This suggests that carbon addition alone is not sufficient to reduce autotrophic production, at least in an oligotrophic lake dominated by mixotrophic phytoplankton. Larval perch grazing did not produce a classical trophic cascade, but substantially altered the species composition of crustacean zooplankton and ciliate trophic levels. Glucose addition increased the biomass of rotifers, thus potentially increasing energy transfer through the heterotrophic pathway, but only when fish were absent. This study illustrates that changes in community structure due to selective feeding by top‐predators can determine the influence of bottom–up carbon subsidies.     

Size-dependent changes in phytoplankton C and N uptake in the dynamic mixed layer of Lake Biwa

1. Short-term (days) hydrodynamic effects of wind-induced mixing on phytoplankton size structure, and C and N uptake characteristics, were studied in the surface mixed layer (epilimnion) of Lake Biwa (North Basin), before and during a period of high winds (typhoons). 2. The latter period was characterized by two major typhoon events associated with deepening of the seasonal thermocline, reduced water column stability, decreased underwater irradiance and increased dissolved reactive N and particulate P. 3. Nutrient concentrations, seston C/N ratios, and uptake rates indicated that phytoplankton biomass and production were limited by P and not N throughout the study. Higher C- and N-based productivity during the typhoon period than before reflected the increased phytoplankton biomass and higher specific uptake rates due to increased nutrient supply. 4. Changes in the size-structure of phytoplankton (< 2 and > 2 μm) were associated with variations in the stratification and mixing regime. When vertical stability was high (before the typhoons) concentrations of > 2 μm biomass (chlorophyll a, particulate organic C and N) were higher at the bottom of the mixed layer than at the surface whereas, when stability of the mixed layer was low (the typhoon period), the contribution of picoplankton (< 2 μm) to total Chl a increased at the surface and decreased at the bottom following the first high winds. 5. Photoadaptive adjustments of the phytoplankton provided further evidence of hydrodynamic control. The lower intracellular Chl a concentrations and C and N uptake efficiencies in the < 2 μm fraction suggest that they experienced, on average, higher irradiance than the larger cells because of their lower sinking rates. During the stability period, picoplankton exhibited higher photosynthetic efficiencies at the bottom of the mixed layer than at the surface. Such differences disappeared during the typhoon period indicating that the mixing rate was then probably higher than the photoacclimation rate in the small size fraction. 6. The present results stress the highly transient nature of biological homogeneity in the surface mixed layer of the lake.     

Diatoms shape the biogeography of heterotrophic prokaryotes in early spring in the Southern Ocean

The interplay among microorganisms profoundly impacts biogeochemical cycles in the ocean. Culture-based work has illustrated the diversity of diatom–prokaryote interactions, but the question of whether these associations can affect the spatial distribution of microbial communities is open. Here, we investigated the relationship between assemblages of diatoms and of heterotrophic prokaryotes in surface waters of the Indian sector of the Southern Ocean in early spring. The community composition of diatoms and that of total and active prokaryotes were different among the major ocean zones investigated. We found significant relationships between compositional changes of diatoms and of prokaryotes. In contrast, spatial changes in the prokaryotic community composition were not related to geographic distance and to environmental parameters when the effect of diatoms was accounted for. Diatoms explained 30% of the variance in both the total and the active prokaryotic community composition in early spring in the Southern Ocean. Using co-occurrence analyses, we identified a large number of highly significant correlations between abundant diatom species and prokaryotic taxa. Our results show that key diatom species of the Southern Ocean are each associated with a distinct prokaryotic community, suggesting that diatom assemblages contribute to shaping the habitat type for heterotrophic prokaryotes.     

Purple Sulfur Bacteria Control the Growth of Aerobic Heterotrophic Bacterioplankton in a Meromictic Salt Lake

In meromictic Mahoney Lake, British Columbia, Canada, the heterotrophic bacterial production in the mixolimnion exceeded concomitant primary production by a factor of 7. Bacterial growth rates were correlated neither to primary production nor to the amount of chlorophyll a. Both results indicate an uncoupling of bacteria and phytoplankton. In the chemocline of the lake, an extremely dense population of the purple sulfur bacterium Amoebobacter purpureus is present year round. We investigated whether anoxygenic phototrophs are significant for the growth of aerobic bacterioplankton in the overlaying water. Bacterial growth rates in the mixolimnion were limited by inorganic phosphorus or nitrogen most of the time, and the biomass of heterotrophic bacteria did not increase until, in autumn, 86% of the cells of A. purpureus appeared in the mixolimnion because of their reduced buoyant density. The increase in heterotrophic bacterial biomass, soluble phosphorus concentrations below the detection limit, and an extraordinarily high activity of alkaline phosphatase in the mixolimnion indicate a rapid liberation of organically bound phosphorus from A. purpureus cells accompanied by a simultaneous incorporation into heterotrophic bacterioplankton. High concentrations of allochthonously derived dissolved organic carbon (mean, 60 mg of C(middot)liter(sup-1)) were measured in the lake water. In Mahoney Lake, liberation of phosphorus from upwelling purple sulfur bacteria and degradation of allochthonous dissolved organic carbon as an additional carbon source render heterotrophic bacterial production largely independent of the photosynthesis of phytoplankton. A recycling of inorganic nutrients via phototrophic bacteria also appears to be relevant in other lakes with anoxic bottom waters.