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.     

Abundance and distribution of nanoplankton in the epipelagic subtropical/tropical open Atlantic Ocean

The objective of this research was to quantify the abundanceand distribution of nanoplanktonic particles available to frequentlyoccurring juvenile and adult planktonic copepods in the epipelagialof the open subtropical and tropical ocean. We chose locationsoff Puerto Rico and off the Bahamas to achieve our purpose.The hydrography for all stations revealed a thin upper mixedlayer, a thermocline with varying temperature decreases, salinitiesapproaching and surpassing 37     

Microbial control of phosphate in the nutrient-depleted North Atlantic subtropical gyre

Little is known about the dynamics of dissolved phosphate in oligotrophic areas of the world's oceans, where concentrations are typically in the nanomolar range. Here, we have budgeted phosphate uptake by the dominant microbial groups in order to assess the effect of the microbial control of this depleted nutrient in the North Atlantic gyre. Low concentrations (2.2 +/- 1.2 nM) and rapid microbial uptake (2.1 +/- 2.4 nM day(-1)) of bioavailable phosphate were repeatedly determined in surface waters of the North Atlantic oligotrophic gyre during spring and autumn research cruises, using a radiotracer dilution bioassay technique. Upper estimates of the concentration of bioavailable phosphate were 7-55% of the dissolved mineral phosphate suggesting that a considerable part of the chemically measured nanomolar phosphate was in a form unavailable for direct microbial uptake. A 1:1 relationship (r(2) = 0.96, P < 0.0001) was observed between the bioavailable total phosphate uptake and the phosphate uptake of all the flow sorted bacterioplankton cells, demonstrating that bacterioplankton were the main consumers of phosphate. Within the bacterioplankton a group of heterotrophic bacteria and Prochlorococcus phototrophic cyanobacteria, were the two major competing groups for bioavailable phosphate. These heterotrophic bacteria had low nucleic acid content and 60% of them comprised of SAR11 clade cells based on the results of fluorescence in situ hybridization. Each of the two competing bacterial groups was responsible for an average of 45% of the phosphate uptake, while Synechococcus cyanobacteria (7%) and picoplanktonic algae (0.3%) played minor roles in direct phosphate uptake. We have demonstrated that phosphate uptake in the oligotrophic gyre is rapid and dominated by two bacterial groups rather than by algae.     

Abundance and single-cell activity of heterotrophic bacterial groups in the western Arctic Ocean in summer and winter

Environmental conditions in the western Arctic Ocean range from constant light and nutrient depletion in summer to complete darkness and sea ice cover in winter. This seasonal environmental variation is likely to have an effect on the use of dissolved organic matter (DOM) by heterotrophic bacteria in surface water. However, this effect is not well studied and we know little about the activity of specific bacterial clades in the surface oceans. The use of DOM by three bacterial subgroups in both winter and summer was examined by microautoradiography combined with fluorescence in situ hybridization. We found selective use of substrates by these groups, although the abundances of Ant4D3 (Antarctic Gammaproteobacteria), Polaribacter (Bacteroidetes), and SAR11 (Alphaproteobacteria) were not different between summer and winter in the Beaufort and Chukchi Seas. The number of cells taking up glucose within all three bacterial groups decreased significantly from summer to winter, while the percentage of cells using leucine did not show a clear pattern between seasons. The uptake of the amino acid mix increased substantially from summer to winter by the Ant4D3 group, although such a large increase in uptake was not seen for the other two groups. Use of glucose by bacteria, but not use of leucine or the amino acid mix, related strongly to inorganic nutrients, chlorophyll a, and other environmental factors. Our results suggest a switch in use of dissolved organic substrates from summer to winter and that the three phylogenetic subgroups examined fill different niches in DOM use in the two seasons.     

Comparison of phosphate uptake rates by the smallest plastidic and aplastidic protists in the North Atlantic subtropical gyre

The smallest phototrophic protists (<3 μm) are important primary producers in oligotrophic subtropical gyres - the Earth's largest ecosystems. In order to elucidate how these protists meet their inorganic nutrient requirements, we compared the phosphate uptake rates of plastidic and aplastidic protists in the phosphate-depleted subtropical and tropical North Atlantic (4-29°N) using a combination of radiotracers and flow cytometric sorting on two Atlantic Meridional Transect cruises. Plastidic protists were divided into two groups according to their size (<2 and 2-3 μm). Both groups of plastidic protists showed higher phosphate uptake rates per cell than the aplastidic protists. Although the phosphate uptake rates of protist cells were on average seven times (P<0.001) higher than those of bacterioplankton, the biomass-specific phosphate uptake rates of protists were one fourth to one twentieth of an average bacterioplankton cell. The unsustainably low biomass-specific phosphate uptake by both plastidic and aplastidic protists suggests the existence of a common alternative means of phosphorus acquisition - predation on phosphorus-rich bacterioplankton cells.     

Abundant proteorhodopsin genes in the North Atlantic Ocean

Proteorhodopsin (PR) is a light-driven proton pump that has been found in a variety of marine bacteria, including Pelagibacter ubique , a member of the ubiquitous SAR11 clade. The goals of this study were to explore the diversity of PR genes and to estimate their abundance in the North Atlantic Ocean using quantitative polymerase chain reaction (QPCR). We found that PR genes in the western portion of the Sargasso Sea could be grouped into 27 clusters, but five clades had the most sequences. Sets of specific QPCR primers were designed to examine the abundance of PR genes in the following four of the five clades: SAR11 ( P. ubique and other SAR11 Alphaproteobacteria ), BACRED17H8 ( Alphaproteobacteria ), HOT2C01 ( Alphaproteobacteria ) and an uncultured subgroup of the Flavobacteria . Two groups (SAR11 and HOT2C01) dominated PR gene abundance in oligotrophic waters, but were significantly less abundant in nutrient- and chlorophyll-rich waters. The other two groups (BACRED17H8 and Flavobacteria subgroup NASB) were less abundant in all waters. Together, these four PR gene types were found in 50% of all bacteria in the Sargasso Sea. We found a significant negative correlation between total PR gene abundance and nutrients and chlorophyll but no significant correlation with light intensity for three of the four PR types in the depth profiles north of the Sargasso Sea. Our data suggest that PR is common in the North Atlantic Ocean, especially in SAR11 bacteria and another marine alphaproteobacterial group (HOT2C01), and that these PR-bearing bacteria are most abundant in oligotrophic waters.     

Deep-sea pelagic ichthyonekton diversity in the Atlantic Ocean and the adjacent sector of the Southern Ocean

Aim Deep‐sea pelagic diversity is poorly understood. Local (SL) and regional (SR) ichthyonekton species richness are presented and analysed with respect to local and regional environmental factors, and biogeographical processes. Location Sixty‐six stations from the Atlantic Ocean and adjacent sector of the Southern Ocean, 65° N to 57° S. Methods Estimation of SL by means of rarefaction. Stepwise evaluation of SL and SR relationships by means of the second‐order corrected Akaike information criterion (AICc) after locally weighted scatterplot smoothing (LOESS) and linear fitting, analysis of saturation effects by means of slopes of species accumulation curves (log–log plots). Results Latitudinal gradients were present for SL and SR, and were asymmetric between the Northern and Southern hemispheres. Relatively low species richness was encountered for the Southern Ocean. Asymmetry at the regional level by means of higher SR was attributed to area effects in the South Atlantic. Log–log plots indicated saturation of local assemblages and dependence on environmental factors. SL was related to productivity; this relationship was hump‐shaped. SR was positively related to area size and negatively to seasonality of production. Biogeographical effects were indicated in that SR peaks coincided with overlap zones of boreal and tropical faunas as a consequence of historical faunal exchange processes. Main conclusions The stepwise approach allowed for distinction between effects of area size, productivity and biogeographical processes on diversity at local and regional scales. Productivity in particular is important in two ways. At the local scale, the link of productivity to SL is explained by a successional‐functional hypothesis of resource utilization, whereas the seasonality effect for SR reinforces the hypothesis of dependence of deep‐sea fishes on seasonality of production through changes of life‐history traits. The causes of low Antarctic faunal diversity remained unresolved.     

Metabolic diversity of heterotrophic bacterioplankton over winter and spring in the coastal Arctic Ocean

Metabolic diversity of heterotrophic bacterioplankton was tracked from early winter through spring with Biolog Ecoplates under the seasonally ice covered arctic shelf in the Canadian Arctic (Franklin Bay, Beaufort Sea). Samples were taken every 6 days from December 2003 to May 2004 at the surface, the halocline where a temperature inversion occurs, and at 200 m, close to the bottom. Despite the low nutrient levels and low chlorophyll a , suggesting oligotrophy in the winter surface waters, the number of substrates used (NSU) was greater than in spring, when chlorophyll a concentrations increased. Denaturing gradient gel electrophorisis analysis also indicated that the winter and spring bacterial communities were phylogenetically distinct, with several new bands appearing in spring. In spring, the bacterial community would have access to the freshly produced organic carbon from the early phytoplankton bloom and the growth of rapidly growing specialist phenotypes would be favoured. In contrast, in winter bacterioplankton consumed more complex organic matter originated during the previous year's phytoplankton production. At the other depths we tested the NSU was similar to that for the winter surface, with no seasonal pattern. Instead, bacterioplankton metabolism seemed to be influenced by resuspension, advection, and sedimentation events that contributed organic matter that enhanced bacterial metabolism.     

Diversity and distribution of single-stranded DNA phages in the North Atlantic Ocean

Knowledge of marine phages is highly biased toward double-stranded DNA (dsDNA) phages; however, recent metagenomic surveys have also identified single-stranded DNA (ssDNA) phages in the oceans. Here, we describe two complete ssDNA phage genomes that were reconstructed from a viral metagenome from 80 m depth at the Bermuda Atlantic Time-series Study (BATS) site in the northwestern Sargasso Sea and examine their spatial and temporal distributions. Both genomes (SARssφ1 and SARssφ2) exhibited similarity to known phages of the Microviridae family in terms of size, GC content, genome organization and protein sequence. PCR amplification of the replication initiation protein (Rep) gene revealed narrow and distinct depth distributions for the newly described ssDNA phages within the upper 200 m of the water column at the BATS site. Comparison of Rep gene sequences obtained from the BATS site over time revealed changes in the diversity of ssDNA phages over monthly time scales, although some nearly identical sequences were recovered from samples collected 4 years apart. Examination of ssDNA phage diversity along transects through the North Atlantic Ocean revealed a positive correlation between genetic distance and geographic distance between sampling sites. Together, the data suggest fundamental differences between the distribution of these ssDNA phages and the distribution of known marine dsDNA phages, possibly because of differences in host range, host distribution, virion stability, or viral evolution mechanisms and rates. Future work needs to elucidate the host ranges for oceanic ssDNA phages and determine their ecological roles in the marine ecosystem.     

Diversity of Diazotrophic Unicellular Cyanobacteria in the Tropical North Atlantic Ocean

We present data on the genetic diversity and phylogenetic affinities of N₂-fixing unicellular cyanobacteria in the plankton of the tropical North Atlantic Ocean. Our dinitrogenase gene (nifH) sequences grouped together with a group of cyanobacteria from the subtropical North Pacific; another subtropical North Pacific group was only distantly related. Most of the 16S ribosomal DNA sequences from our tropical North Atlantic samples were closely allied with sequences from a symbiont of the diatom Climacodium frauenfeldianum. These findings suggest a complex pattern of evolutionary and ecological divergence among unicellular cyanobacteria within and between ocean basins.     

Description of phytoplankton and nutrient in spring in the western North Atlantic Ocean

Abundance of diatoms in the northern Sargasso Sea in springhas been observed in a 4-year period between 1981 and 1985 andthen again in 1987. A theory is presented that if stratificationblocks get-through of deep nutrients to the surface, as in thesouthern Sargasso, then in-situ cycling via non-diatoms andammonia dominates. The theory is that if in-situ cycling doesnot dominate, as northward, then stratification does not blockget-through of deep nutrients to the surface, which, in combinationwith the marked growth capacity of diatoms, produce an abundanceof cells. This abundance consumes nitrate, phosphate and silicateat the surface and ultimately this nutrient-depleted water arrivesin the southern Sargasso via the gyre circulation. Thus thenutrient-depleted water of the western Atlantic gyre is consideredto be primarily achieved only by diatoms, in spring and in thenorthern Sargasso. A transect during April 13–16, 1985,north from the Virgin Islands along 63°–64°W showsa great change in the diatom species plus Emiliania huxleyifrom negligible concentrations in the southern half of the sectionto large concentrations in the northern half. This unevennessof distribution can be rephrased by indicating that these speciesare narrow-niched with regard to nutrient availability. Theother species, coccolithophores and dinoflagellates, are evenlydistributed at low concentrations south-north (though some diminishslightly) and so they are broad-niched with respect to nutrient.High species diversity depends on broadness of nutrient nicheand equilibrium conditions of hydrographically stable water(i), implying that when non-equilibrium conditions occur (ii),when niche breadth is less (iii) or both (iv), then diversityis less marked, (i) is confirmed by broad-niched species southward,(ii) is confirmed by broad-niched species northward, (iii) isconfirmed by narrow-niched species southward and (iv) by narrow-nichedspecies northward.     

Abundance and biogeography of tintinnids (Ciliophora) and associated microzooplankton in the Southwestern Atlantic Ocean

Absolute abundances of foraminifers, polycystine and phaeodarian radiolarians, tintinnids, pteropods and early crustacean larvae and moults were assessed in a collection of 57 vertically stratified (0-100 m) net microplankton samples from 22 stations located between 34 and 58S (along 51-56°W), covered on 8-16 November 1994. Tintinnids were identified to species and measured in order to estimate their biomass from biovolume to carbon conversions. The distribution of the microzooplanktonic groups assessed was irregular and patchy, both geographically and vertically, and their abundances were characteristic of oceanic low to medium productivity environments. Tintinnid biomass was also generally low (0.05-0.40 g Cl^-1). With the exception of the tintinnids, associations between microzooplanktonic numbers and chlorophyll a were generally loose. Eighty-eight tintinnid taxa were recorded, yet only five accounted for 53% of the specimens identified. Multivariate (cluster) analysis of tintinnid specific distribution patterns clearly showed several distinct zones. From north to south, these are: Transition Zone (TZ), with three subzones, TZ north (34°S-38°S), TZ central (39°44S-44°S) and TZ south (46°S); Subantarctic Zone (SZ; 48-55°S); Polar Front Zone (55°30S); Antarctic Zone (AZ; 58-59°36S). Each of these was characterized by distinct tintinnid assemblages, abundance and biomass. With few exceptions, tintinnid cells were fairly evenly distributed throughout the upper 50 m. Taxonomic composition usually changed little with depth. Mean population depths were calculated for a subset of 35 tintinnids; 29 of these dwell preferably above 40 m. The spatial distribution of tintinnid species richness showed a more or less gradual decrease from north to south. Specific diversity and equitability generally increased with depth, and were higher in antarctic waters than the southern transitional and subantarctic ones; this trend is tentatively attributed to higher water column vertical stability south of the Polar Front.      

Pollen distribution in the bottom sediments of the western North Atlantic Ocean

An extensive survey of pollen distribution in surficial sediments from the slope and rise between 29° and 72°N, shows patterns which reflect the distribution of vegetation in eastern North America and the effects of aeolian, fluvial, and marine sedimentation. The results of the analysis of 94 core tops show that pollen concentration ranges from 23,000 to 10 grains per gram, and decreases offshore with distance from vegetation sources and northward in relation to lower vegetation density. Maximum concentration occurs in organic-rich, fine-grained lutum deposited off rivers draining the southeastern United States. Low pollen concentrations are associated with high-energy environments, such as the Western Boundary Undercurrent and the Labrador Current. Percentages of dominant pollen types generally correspond with onshore distribution of pollen and vegetation. Oak and other deciduous pollen types are the most abundant in marine sediments off the deciduous forests of eastern United States. Hemlock pollen is essentially restricted to sediments off the hemlock-oak forests of northeastern U.S. and southeastern Canada, and spruce pollen is most abundant adjacent boreal forests and forest-tundra formations. The relative increase in pine pollen seaward reflects the selective effects of marine transport. Synthesis of pollen data by Q-mode factor analysis shows that major vegetation formations of eastern North America produce distinctive, regional marine pollen assemblages.     

Orientation behaviour of leatherback sea turtles within the North Atlantic subtropical gyre

Leatherback sea turtles (Dermochelys coriacea) travel thousands of kilometres between temperate feeding and tropical breeding/over-wintering grounds, with adult turtles able to pinpoint specific nesting beaches after multi-year absences. Their extensive migrations often occur in oceanic habitat where limited known sensory information is available to aid in orientation. Here, we examined the migratory orientation of adult male, adult female and subadult leatherbacks during their open-ocean movements within the North Atlantic subtropical gyre by analysing satellite-derived tracks from fifteen individuals over a 2-year period. To determine the turtles'' true headings, we corrected the reconstructed tracks for current drift and found negligible differences between current-corrected and observed tracks within the gyre. Individual leatherback headings were remarkably consistent throughout the subtropical gyre, with turtles significantly oriented to the south-southeast. Adult leatherbacks of both sexes maintained similar mean headings and showed greater orientation precision overall. The consistent headings maintained by adult and subadult leatherbacks within the gyre suggest use of a common compass sense.     

Nitrogen and phosphorus budgets of the North Atlantic Ocean and its watershed

Anthropogenic food and energy production extensively mobilize reactive nitrogen (N) in the watershed of the North Atlantic Ocean (NAO). There is wide spread N distribution by both hydrologic and atmospheric processes within the watershed of the NAO, resulting in reactive N accumulation in terrestrial systems. Net denitrification in most estuaries and continental shelves exceeds the amount of N supplied to the shelves by rivers and requires a supply of nitrate from the open ocean. Thus riverine N is only transported to the open ocean in a few areas with the flow from a few major rivers (e.g., Amazon). Atmospheric N deposition to the open ocean has increased and may increase the productivity of the surface ocean. In addition, as a consequence of increased Fe deposition to the open ocean (due in part to anthropogenic processes), the rate of biological N-fixation may have increased resulting in N accumulation in the ocean. Phosphorus (P) is also mobilized by anthropogenic processes (primarily food production). Relative to N, more of the P is transported across the shelf to the open ocean from both estuaries and major rivers. There are several consequences of the increased availability of N and P that are unique to each element. However, the control on primary productivity in both coastal and open ocean ecosystems is dependent on a complex and poorly understood interaction between N and P mobilization and availability.     

The distribution of seaweed floras in the tropical and subtropical Atlantic Ocean: a quantitative approach

Records of nearly 1500 species of marine algae present in over 80 countries, states, and islands of the tropical and subtropical Atlantic were analyzed to ascertain the relationships between their floras. Association analysis was unsatisfactory, cluster analysis was better, but, of the classificatory methods, indicator species analysis was best. However, the most meaningful results were obtained by ordination using reciprocal averaging. This method demonstrates that after the separation of the marine flora of South-West Africa (Namibia), two clearly defined groupings representing the eastern Atlantic and the western Atlantic are apparent. On the western side, the floras, apart from that of Uruguay, are a relatively close-knit group, except that the northern coast of the Gulf of Mexico together with the southern Atlantic States of the U.S.A. are distinct from the southern Gulf of Mexico and Caribbean groups. On the eastern side of the Atlantic the floras are less closely related. A well-marked tropical group extends from Gambia to Cameroun but does not include the Gulf of Guinea islands which form another group with Angola. This latter group is transitional to the colder water floras lying northwards of Senegal and south of Angola.     

Diversity and abundance of glycosyl hydrolase family 5 in the North Atlantic Ocean

The diversity and abundance of glycosyl hydrolase family 5 (GH5) were studied in the North Atlantic Ocean. This family was chosen because of the large number of available sequences from cultured bacteria, the variety of substrates it targets, and the high number of similar sequences in the Sargasso Sea environmental genome database. Three clone libraries of a GH5 subcluster were constructed from the Mid-Atlantic Bight and the eastern and western North Atlantic Ocean. The two North Atlantic Ocean libraries did not differ from each other but both were significantly less diverse than the Mid-Atlantic Bight library. The abundance of GH5 genes estimated by quantitative PCR was positively correlated with chlorophyll concentrations in the eastern part of a transect from Fort Pierce, Florida, to the Azores and in a depth profile, suggesting that the supply of labile organic material selects for GH5-bearing bacteria in these waters. However, the data suggest that only <1% of all bacteria harbor the GH5 subcluster. These and other data suggest that the hydrolysis of polysaccharides requires complicated multi-enzyme systems.     

Latitudinal variation in virus-induced mortality of phytoplankton across the North Atlantic Ocean

Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeochemical cycles in the ocean. Yet, little is known about the biogeographical distribution of viral lysis rates across the global ocean. To address this, we investigated phytoplankton group-specific viral lysis rates along a latitudinal gradient within the North Atlantic Ocean. The data show large-scale distribution patterns of different virus groups across the North Atlantic that are associated with the biogeographical distributions of their potential microbial hosts. Average virus-mediated lysis rates of the picocyanobacteria Prochlorococcus and Synechococcus were lower than those of the picoeukaryotic and nanoeukaryotic phytoplankton (that is, 0.14 per day compared with 0.19 and 0.23 per day, respectively). Total phytoplankton mortality (virus plus grazer-mediated) was comparable to the gross growth rate, demonstrating high turnover rates of phytoplankton populations. Virus-induced mortality was an important loss process at low and mid latitudes, whereas phytoplankton mortality was dominated by microzooplankton grazing at higher latitudes (>56°N). This shift from a viral-lysis-dominated to a grazing-dominated phytoplankton community was associated with a decrease in temperature and salinity, and the decrease in viral lysis rates was also associated with increased vertical mixing at higher latitudes. Ocean-climate models predict that surface warming will lead to an expansion of the stratified and oligotrophic regions of the world''s oceans. Our findings suggest that these future shifts in the regional climate of the ocean surface layer are likely to increase the contribution of viral lysis to phytoplankton mortality in the higher-latitude waters of the North Atlantic, which may potentially reduce transfer of matter and energy up the food chain and thus affect the capacity of the northern North Atlantic to act as a long-term sink for CO₂.