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.     

Resource-Limited Heterotrophic Prokaryote Production and Its Potential Environmental Impact Associated with Mn Nodule Exploitation in the Northeast Equatorial Pacific

Shipboard enrichment incubation experiments were performed to elucidate the limiting resources for heterotrophic prokaryotic production and to discuss the potential impact of bottom water and sediment discharges in relation to manganese (Mn) nodule exploitation on the heterotrophic prokaryotes in the oligotrophic northeast equatorial Pacific. Compared to an unamended control, the production of heterotrophic prokaryotes increased 25-fold in water samples supplemented with amino acids (i.e., organic carbon plus nitrogen), whereas the production increased five and two times, respectively, in samples supplemented with either glucose or ammonium alone. These results indicate that heterotrophic prokaryote production in the northeast equatorial Pacific was co-limited by the availability of dissolved organic carbon and inorganic nitrogen. In samples from the nutrient-depleted surface mixed layer (10-m depth), the addition of a slurry of bottom water and sediment doubled heterotrophic prokaryote production compared to an unamended control, whereas sonicating the slurry prior to addition quadrupled the production rate. However, little difference was observed between an unamended control and slurry-amended samples in the subsurface chlorophyll a (Chl a) maximum (SCM) layer. Thus, the impact of slurry discharge is more significant at the nutrient-depleted surface mixed layer than at the high-nutrient SCM layer. The greatly enhanced prokaryote production resulting from the addition of sonicated slurry further suggests that dissociated organic carbon may directly stimulate heterotrophic prokaryote production in the surface mixed layer. Overall, the results suggest that the surface discharge of bottom water and sediments during manganese nodule exploitation could have a significant environmental impact on the production of heterotrophic prokaryotes that are currently resource limited.     

Prokaryotic extracellular enzymatic activity in relation to biomass production and respiration in the meso- and bathypelagic waters of the (sub)tropical Atlantic

Prokaryotic extracellular enzymatic activity, abundance, heterotrophic production and respiration were determined in the meso- and bathypelagic (sub)tropical North Atlantic. While prokaryotic heterotrophic production (PHP) decreased from the lower euphotic layer to the bathypelagic waters by two orders of magnitude, prokaryotic abundance and cell-specific PHP decreased only by one order of magnitude. In contrast to cell-specific PHP, cell-specific extracellular enzymatic activity (alpha- and beta-glucosidase, leucine aminopeptidase, alkaline phosphatase) increased with depth as did cell-specific respiration rates. Cell-specific alkaline phosphatase activity increased from the intermediate water masses to the deep waters up to fivefold. Phosphate concentrations, however, varied only by a factor of two between the different water masses, indicating that phosphatase activity is not related to phosphate availability in the deep waters. Generally, cell-specific extracellular enzymatic activities were inversely related to cell-specific prokaryotic leucine incorporation. Thus, it is apparent that the utilization of deep ocean organic matter is linked to higher cell-specific extracellular enzymatic activity and respiration and lower cell-specific PHP than in surface waters.     

Plankton metabolism in the Greenland Sea during the polar summer of 2007

The polar summer metabolism of the planktonic communities in the Greenland Sea was surveyed in July 2007. Planktonic metabolism showed great variability across the studied area, with on average, higher metabolic rates in the Fram Strait-Svalbard region than along the Greenland Current. A significant fraction (47%) of the planktonic communities in the Fram Strait-Svalbard region were net heterotrophic, suggesting that increased respiration rates with further warming may lead the planktonic communities at this region to act as net CO₂ sources. The thresholds gross primary production for metabolic balance (i.e., gross primary production = community respiration) was much higher in the European sector of the Arctic than reported for the Southern Ocean, suggesting that heterotrophic metabolism is more prevalent in the European sector of the Arctic than in the Southern Ocean, indicating high allochthonous inputs in the Arctic region.     

Spatial and seasonal changes of dissolved and particulate organic C in the North Sea

Sampling of the central region of the North Sea was carried out to study the spatial and seasonal changes of dissolved and particulate organic C (DOC and POC, respectively). The surface waters were collected during four cruises over a year (Autumn 2004–Summer 2005). DOC and POC concentrations were measured using high temperature catalytic oxidation methods. The surface water concentrations of DOC and POC were spatially and temporally variable. There were significantly different concentrations of DOC and POC between the inshore and offshore waters in winter and summer only, with no clear trend in autumn and spring. Highest mean concentrations of DOC were measured in spring with lower and similar mean concentrations for other seasons. POC showed a clear seasonal cycle throughout the year with highest surface mean concentrations found in autumn and spring, but lowest in winter and summer. The DOC distributions during autumn and spring were strongly correlated with chlorophyll suggesting extracellular release from phytoplankton was an important DOC source during these two seasons. The lower concentrations of DOC in summer were probably due to the heterotrophic uptake of labile DOC. The seasonal changes in the C:N molar ratios of surface DOM (dissolved organic matter) resulted in higher mean C:N molar ratios in spring and lower ratios in winter. These high ratios may indicate nutrient limitation of heterotrophic uptake immediately after the spring bloom. There is limited data available for DOC cycling in these productive shelf seas and these results show that DOC is a major component of the C cycle with partial decoupling of the DOC and DON cycling in the central North Sea during the spring bloom. Handling editor: Luigi Naselli-Flores     

Seasonal Depth-Related Gradients in Virioplankton: Lytic Activity and Comparison with Protistan Grazing Potential in Lake Pavin (France)

This study presents an original depth-related survey of virioplankton lytic activity in relation to prokaryotic production and potential protistan bacterivory in the deep (Z(max) = 92 m) meromictic volcanic Lake Pavin (Massif Central, France). The sampling strategy was designed to be representative of the physico-chemical gradients of the water column of the lake, and of the seasonal variability as well, i.e. 12 different depths sampled in triplicates from April to December 2005. In the space, viral lytic activity estimated from the frequency of visibly infected prokaryotic cells and from burst size over the study period generally decreased with depth. This was viewed as a paradox compared to the abundances of viruses and prokaryotes and to the prokaryotic production which increased with depth. The seasonal variability in viral lytic activity was correlated with prokaryotic variables (abundance and production) in the deepest waters, i.e. from the hypolimnion downwards. Compared to previous studies known from the mixolimnion, we conclude that the deep waters in Lake Pavin represent an exclusive environment for heterotrophic prokaryotes whose seasonal activity offers an optimal and unique resource for thriving viral communities, some of which may be typical, endemic to the ambient dark, cold and stable deep water masses. Overall, the main findings in the present study get well around a previous statement that the ecology of the deepest waters of Lake Pavin is essentially driven by the dark viral loop (dissolved organic matter-prokaryotes-viruses) processes, which can sequester organic matters and nutrients for a long-lived turnover time. This is in agreement with recent demonstrations from marine systems that meso- and bathypelagic waters are optimal environments for viral survival and proliferation.     

Psychrophilic versus psychrotolerant bacteria--occurrence and significance in polar and temperate marine habitats.

The numerical dominance and ecological role of psychrophilic bacteria in bottom sediments, sea ice, surface water and melt pools of the polar oceans were investigated using isolates, colony forming units (CFU) and metabolic activities. All sediment samples of the Southern Ocean studied showed a clear numerical dominance of cold-loving bacteria. In Arctic sediments underlying the influence of cold polar water bodies psychrophiles prevailed also but they were less dominant in sediments influenced by the warm Atlantic Water. A predominance of psychrophiles was further found in consolidated Antarctic sea ice as well as in multiyear Arctic sea ice and in melt pools on top of Arctic ice floes. A less uniform adaptation response was, however, met in polar surface waters. In the very northern part of the Fram Strait (Arctic Ocean) we found bacterial counts and activities at 1 degree C exceeding those at 22 degrees C. In surface water of the Weddell Sea (Southern Ocean) psychrophiles also dominated numerically in early autumn but the dominance declined obviously with the onset of winter-water and a decrease of chlorphyll a. Otherwise in surface water of the Southern Ocean CFUs were higher at 22 degrees C than at 1 degree C while activities were vice versa indicating at least a functional dominance of psychrophiles. Even in the temperate sediments of the German Bight true psychrophiles were present and a clear shift towards cold adapted communities in winter observed. Among the polar bacteria a more pronounced cold adaptation of Antarctic in comparison with Arctic isolates was obtained. The results and literature data indicate that stenothermic cold adapted bacteria play a significant role in the global marine environment. On the basis of the temperature response of our isolates from different habitats it is suggested to expand the definition of Morita in order to meet the cold adaptation strategies of the bacteria in the various cold habitats.     

Microzooplankton Growth Rates Examined across a Temperature Gradient in the Barents Sea

Growth rates (µ) of abundant microzooplankton species were examined in field experiments conducted at ambient sea temperatures (−1.8–9.0°C) in the Barents Sea and adjacent waters (70–78.5°N). The maximum species-specific µ of ciliates and athecate dinoflagellates (0.33–1.67 d⁻¹ and 0.52–1.14 d⁻¹, respectively) occurred at temperatures below 5°C and exceeded the µ_max predicted by previously published, laboratory culture-derived equations. The opposite trend was found for thecate dinoflagellates, which grew faster in the warmer Atlantic Ocean water. Mixotrophic ciliates and dinoflagellates grew faster than their heterotrophic counterparts. At sub-zero temperatures, microzooplankton µ_max matched those predicted for phytoplankton by temperature-dependent growth equations. These results indicate that microzooplankton protists may be as adapted to extreme Arctic conditions as their algal prey.     

Experimental evaluation of planktonic respiration response to warming in the European Arctic Sector

The Arctic Ocean is the region on Earth supporting the steepest warming rate and is also particularly vulnerable due to the vanishing ice cover. Intense warming in the Arctic has strong implications for biological activity and the functioning of an Arctic Ocean deprived of ice cover in summer. We evaluated the impact of increasing temperature on respiration rates of surface marine planktonic communities in the European Arctic sector, a property constraining the future role of the Arctic Ocean in the CO₂ balance of the atmosphere. We performed experiments under four different temperature elevation regimes (in situ, +2, +4 and +6°C above the temperature of the sampled water) during cruises conducted in the Fram Strait region and off Svalbard during late fall–early winter, spring and summer. During late fall–early winter, where only three different temperatures were used, no response to warming was observed, whereas respiration rates increased in response to warming in spring and summer, although with variable strength.     

The role of dissolved organic matter bioavailability in promoting phytoplankton blooms in Florida Bay

The clear, shallow, oligotrophic waters of Florida Bay are characterized by low phytoplankton biomass, yet periodic cyanobacteria and diatom blooms do occur. We hypothesized that allochthonous dissolved organic matter (DOM) was providing a subsidy to the system in the form of bound nutrients. Water from four bay sites was incubated under natural light and dark conditions with enrichments of either DOM ( > 1 kD, 2×DOM) or inorganic nutrients (N+P). Samples were analyzed for bacterial numbers, bacterial production, phytoplankton biomass, phytoplankton community structure, and production, nutrients, and alkaline phosphatase (AP) activity. The influence of 2×DOM enrichment on phytoplankton biomass developed slowly during the incubations and was relatively small compared to nutrient additions. Inorganic nutrient additions resulted in an ephemeral bloom characterized initially as cyanobacterial and brown algae but which changed to dinoflagellate and/or brown algae by day six. The DIN:TP ratio decreased 10-fold in the N+P treatments as the system progressed towards N limitation. This ratio did not change significantly for 2×DOM treatments. In addition, these experiments indicated that both autotrophic and heterotrophic microbial populations in Florida Bay may fluctuate in their limitation by organic and inorganic nutrient availability. Both N+P and 2×DOM enrichments revealed significant and positive response in bioavailability of dissolved organic carbon (BDOC). Potential BDOC ranged from 1.1 to 35.5%, with the most labile forms occurring in Whipray Basin. BDOC at all sites was stimulated by the 2×DOM addition. Except for Duck Key, BDOC at all sites was also stimulated by the addition of N+P. BDOC was lower in the dry season than in the wet season (5.56% vs. 16.86%). This may be explained by the distinct chemical characteristics of the DOM produced at different times of year. Thus, both the heterotrophic and autotrophic microbial communities in Florida Bay are modulated by bioavailability of DOM. This has ramifications for the fate of DOM from the Everglades inputs, implicating DOM bioavailability as a contributing factor in regulating the onset, persistence, and composition of phytoplankton blooms.     

Diazotrophic diversity and distribution in the tropical and subtropical Atlantic Ocean

To understand the structure of marine diazotrophic communities in the tropical and subtropical Atlantic Ocean, the molecular diversity of the nifH gene was studied by nested PCR amplification using degenerate primers, followed by cloning and sequencing. Sequences of nifH genes were amplified from environmental DNA samples collected during three cruises (November-December 2000, March 2002, and October-November 2002) covering an area between 0 to 28.3 degrees N and 56.6 to 18.5 degrees W. A total of 170 unique sequences were recovered from 18 stations and 23 depths. Samples from the November-December 2000 cruise contained both unicellular and filamentous cyanobacterial nifH phylotypes, as well as gamma-proteobacterial and cluster III sequences, so far only reported in the Pacific Ocean. In contrast, samples from the March 2002 cruise contained only phylotypes related to the uncultured group A unicellular cyanobacteria. The October-November 2002 cruise contained both filamentous and unicellular cyanobacterial and gamma-proteobacterial sequences. Several sequences were identical at the nucleotide level to previously described environmental sequences from the Pacific Ocean, including group A sequences. The data suggest a community shift from filamentous cyanobacteria in surface waters to unicellular cyanobacteria and/or heterotrophic bacteria in deeper waters. With one exception, filamentous cyanobacterial nifH sequences were present within temperatures ranging between 26.5 and 30 degrees C and where nitrate was undetectable. In contrast, nonfilamentous nifH sequences were found throughout a broader temperature range, 15 to 30 degrees C, more often in waters with temperature of <26 degrees C, and were sometimes recovered from waters with detectable nitrate concentrations.     

Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Central Arctic Ocean paleoceanographic interpretations based on Late Cenozoic calcareous dinoflagellates

Four calcareous dinoflagellate morphotypes, interpreted to represent a single species, Thoracosphaera arctica n. sp., occur in upper Miocene to Holocene sediments of the central Arctic Ocean. The calcareous fossils generally are rare except during one interval, that of the latest Pliocene to late Pleistocene. During this interval the fossil dinoflagellates were alternately abundant and rare in approximate coordination with greater and lesser times of foraminifera productivity. Dinoflagellates and foraminifera generally are rare during times of accumulation of abundant coarse textured ice-rafted sediment during this interval.The life cycle of the dinoflagellate that produces the Arctic Ocean calcareous fossil is unknown but the fossils probably are either resting cysts, motile vegetative cells or hypnozygotes. A vegetative cell origin better explains the co-occurrence of foraminifera as well as the negative correlation with ice-rafting. This tentative interpretation leads to the conclusion that during the interval 2.0 to 0.7 m.y. b.p., the Arctic Ocean had relatively productive surface waters with very thin or even periodic absence of pack-ice alternating with conditions more like that of the present. During periods before 2.0 m.y. and since 0.7 m.y., the Arctic Ocean probably was ice-covered, with oceanographic conditions similar to that of the modern ocean. These conclusions differ from previous interpretations that the Arctic was continually ice-covered, had no surface productivity or was covered by a giant, Antarctic-like ice-cap.     

Influence of weather on reproductive success of northern fulmars in the Canadian high Arctic

The northern fulmar (Fulmarus glacialis) is a common seabird of the North Atlantic Ocean, with breeding colonies broadly dispersed between 45°N and 80°N. At higher latitudes, breeding fulmars experience extensive sea-ice and presumably snow and low temperatures which do not affect fulmars in the southern part of the breeding range. We studied the relationship between weather and reproductive success of northern fulmars breeding at two colonies in the Canadian high Arctic. Collectively, hatching success, fledging success, and productivity (chicks fledged per egg laid) were similar between our study and results from colonies located south of the Arctic. However, a larger proportion of fulmars at apparently occupied sites (AOS) in high Arctic colonies appeared to forego egg-laying, resulting in lower proportions of chicks fledged per AOS. Extreme inclement weather was the major factor influencing nesting success, resulting in pulses of egg or chick loss during or immediately following major storms, although the mechanism of effects appeared to differ between the two colonies. For Arctic fulmars, the risks of nest failure due to stochastic, deleterious weather events may be offset by the predictable abundance of food supplies during chick-rearing in Arctic waters.     

Cumaceans as Indicators of Atlantic Waters over the Continental Slope of the Arctic Ocean

The material on cumacean fauna of the continental slope of the Arctic Ocean (more than 80 samples) was collected during expeditions with the ships Persei in 1925, 1928, and 1936; Sadko in 1936; and Polarstern in 1990, 1993, and 1995. Examination of the samples (150–900 m depth) revealed 19 species of Cumacea in this region. Three Arcto-Atlantic bathyal species (Hemilamprops uniplicatus, Diastylis echinata, Campylaspis globosa) are markers of the places over the continental slope where Atlantic intermediate waters penetrate into the Arctic Ocean and reach to the bottom. These species are widespread in the North Atlantic (100 to 2882 m depth); on the continental slope of the Arctic Ocean, they occur in a narrow depth range (200 to 900 m) in areas where positive temperatures from 0.09 to 1.78°C and high salinity of 34.75 to 34.87 prevail. The locations of these species is delineated on maps as an interrupted belt extending from northeastern Greenland to the East Siberian Sea and further north up to 79°30 N and 148° E, which corresponds to the route of penetration of Atlantic intermediate waters into the Arctic Ocean.     

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.     

New Production Regulates Export Stoichiometry in the Ocean

The proportion in which carbon and growth-limiting nutrients are exported from the oceans’ productive surface layer to the deep sea is a crucial parameter in models of the biological carbon pump. Based on >400 vertical flux observations of particulate organic carbon (POC) and nitrogen (PON) from the European Arctic Ocean we show the common assumption of constant C:N stoichiometry not to be met. Exported POC:PON ratios exceeded the classical Redfield atomic ratio of 6.625 in the entire region, with the largest deviation in the deep Central Arctic Ocean. In this part the mean exported POC:PON ratio of 9.7 (a:a) implies c. 40% higher carbon export compared to Redfield-based estimates. When spatially integrated, the potential POC export in the European Arctic was 10–30% higher than suggested by calculations based on constant POC:PON ratios. We further demonstrate that the exported POC:PON ratio varies regionally in relation to nitrate-based new production over geographical scales that range from the Arctic to the subtropics, being highest in the least productive oligotrophic Central Arctic Ocean and subtropical gyres. Accounting for variations in export stoichiometry among systems of different productivity will improve the ability of models to resolve regional patterns in carbon export and, hence, the oceans’ contribution to the global carbon cycle will be predicted more accurately.     

Microbial functioning and community structure variability in the mesopelagic and epipelagic waters of the subtropical northeast atlantic ocean

We analyzed the regional distribution of bulk heterotrophic prokaryotic activity (leucine incorporation) and selected single-cell parameters (cell viability and nucleic acid content) as parameters for microbial functioning, as well as bacterial and archaeal community structure in the epipelagic (0 to 200 m) and mesopelagic (200 to 1,000 m) subtropical Northeast Atlantic Ocean. We selectively sampled three contrasting regions covering a wide range of surface productivity and oceanographic properties within the same basin: (i) the eddy field south of the Canary Islands, (ii) the open-ocean NE Atlantic Subtropical Gyre, and (iii) the upwelling filament off Cape Blanc. In the epipelagic waters, a high regional variation in hydrographic parameters and bacterial community structure was detected, accompanied, however, by a low variability in microbial functioning. In contrast, mesopelagic microbial functioning was highly variable between the studied regions despite the homogeneous abiotic conditions found therein. More microbial functioning parameters indicated differences among the three regions within the mesopelagic (i.e., viability of cells, nucleic acid content, cell-specific heterotrophic activity, nanoflagellate abundance, prokaryote-to-nanoflagellate abundance ratio) than within the epipelagic (i.e., bulk activity, nucleic acid content, and nanoflagellate abundance) waters. Our results show that the mesopelagic realm in the Northeast Atlantic is, in terms of microbial activity, more heterogeneous than its epipelagic counterpart, probably linked to mesoscale hydrographical variations.     

Distribution of Roseobacter RCA and SAR11 lineages and distinct bacterial communities from the subtropics to the Southern Ocean

We assessed the composition of the bacterioplankton in the Atlantic sector of the Southern Ocean in austral fall and winter and in New Zealand coastal waters in summer. The various water masses between the subtropics/Agulhas–Benguela boundary region and the Antarctic coastal current exhibited distinct bacterioplankton communities with the highest richness in the polar frontal region, as shown by denaturing gradient gel electrophoresis of 16S rRNA gene fragments. The SAR11 clade and the Roseobacter clade‐affiliated (RCA) cluster were quantified by real‐time quantitative PCR. SAR11 was detected in all samples analysed from subtropical waters to the coastal current and to depths of > 1000 m. In fall and winter, this clade constituted < 3% to 48% and 4–28% of total bacterial 16S rRNA genes respectively, with highest fractions in subtropical to polar frontal regions. The RCA cluster was only present in New Zealand coastal surface waters not exceeding 17°C, in the Agulhas–Benguela boundary region (visited only during the winter cruise), in subantarctic waters and in the Southern Ocean. In fall, this cluster constituted up to 36% of total bacterial 16S rRNA genes with highest fractions in the Antarctic coastal current and outnumbered the SAR11 clade at most stations in the polar frontal region and further south. In winter, the RCA cluster constituted lower proportions than the SAR11 clade and did not exceed 8% of total bacterial 16S rRNA genes. In fall, the RCA cluster exhibited significant positive correlations with latitude and ammonium concentrations and negative correlations with concentrations of nitrate, phosphate, and for near‐surface samples also with chlorophyll a, biomass production of heterotrophic prokaryotes and glucose turnover rates. The findings show that the various water masses between the subtropics and the Antarctic coastal current harbour distinct bacterioplankton communities. They further indicate that the RCA cluster, despite the narrow sequence similarity of > 98% of its 16S rRNA gene, is an abundant component of the heterotrophic bacterioplankton in the Southern Ocean, in particular in its coldest regions.     

Effects of Temperature on Two Psychrophilic Ecotypes of a Heterotrophic Nanoflagellate, Paraphysomonas imperforata

Two different psychrophilic types of the heterotrophic nanoflagellate Paraphysomonas imperforata were isolated from Newfoundland coastal waters and the Arctic Ocean. When fed bacteria without food limitation, both isolates were able to grow at temperatures from -1.8 to 20°C, with maximum growth rates of 3.28 day^-1 at 15°C and 2.28 day^-1 at 12.3°C for the Newfoundland and the Arctic isolates, respectively. Ingestion rates increased with temperature from 14 to 62 bacteria flagellate^-1 h^-1 for the Newfoundland isolate and from 30 to 99 bacteria flagellate^-1 h^-1 for the Arctic isolate. While temperature did not affect cell yields (number of protozoa produced divided by number of bacteria consumed), it affected flagellate sizes. This differential effect of temperature on cell yield and cell size resulted in a changing gross growth efficiency (GGE) in terms of biovolume; colder temperatures favored higher GGEs. The comparison of Q₁₀ values for growth rates and ingestion rates between the isolates shows that the Arctic isolate is better adapted to extremely cold temperature than the Newfoundland isolate. At seawater-freezing temperature (-1.8°C), the estimated maximum growth rates and maximum ingestion rates are 0.81 day^-1 and 30 bacteria flagellate^-1 h^-1 for the Arctic isolate and 0.54 day^-1 and 12 bacteria flagellate^-1 h^-1 for the Newfoundland isolate. Our findings about psychrophilic nanoflagellates fit the general characteristics of cold-water-dwelling organisms: reduced physiological rates and higher GGEs at lower temperatures. Because of the large and persistent differences between the isolates, we conclude that they are ecotypes adapted to specific environmental conditions.