Aphotic Waters Research Articles

Winter Convective Mixing Mediating Coupling of N‐Gain and ‐Loss in the Arabian Sea

AbstractMarine dinitrogen (N2) fixation fuels primary production and thereby influences the Earth's climate. Yet, its geographical distribution and controlling environmental parameters remain debatable. We measured N2 fixation rates from the two spatially and physicochemically contrasting regions of the Arabian Sea during the winter monsoon: (a) the colder and nutrient‐rich waters in the northern region owing to winter convection and (b) the warmer and nutrient‐poor waters in the southern region unaffected by winter convection. We found higher N2 fixation rates at the surface of northern region due to convective mixing driven supply of phosphate (intuitively iron also) from the underlying suboxic waters. N2 fixation was favored by high nutrient concentrations in the euphotic waters, whereas remained unaffected by nutrient availability in the aphotic waters. We conclude that diazotrophs dwelling in the euphotic zone chose phosphate over fixed nitrogen‐poor waters. However, we found that among oligotrophic waters, anticyclonic eddy extremes the barrier of fixed nitrogen supply, and thereby, elevates N2 fixation. While the Arabian Sea loses about 20%–40% of the global ocean fixed nitrogen, we estimate that N2 fixation in the Arabian Sea offsets only up to 42% of its fixed nitrogen‐loss by denitrification, but this offset could be higher if diazotrophic activity is further examined up to the deeper depths of the Arabian Sea.

Preserved particulate organic carbon is likely derived from the subsurface sulfidic photic zone of the Proterozoic Ocean: evidence from a modern, oxygen-deficient lake.

Biological processes in the Proterozoic Ocean are often inferred from modern oxygen-deficient environments (MODEs) or from stable isotopes in preserved sediment. To date, few MODE studies have simultaneously quantified carbon fixation genes and attendant stable isotopic signatures. Consequently, how carbon isotope patterns reflect these pathways has not been thoroughly vetted. Addressing this, we profiled planktonic productivity and quantified carbon fixation pathway genes and associated organic carbon isotope values (δ13 CPOC ) of size-fractionated (0.2-2.7 and >2.7 μm) particulate matter from meromictic Fayetteville Green Lake, NY, USA. The high-O2 Calvin-Benson-Bassham (CBB) gene (cbbL) was most abundant in the <2.7 μm size fraction in shallow oxic and deep hypoxic waters, corresponding with cyanobacterial and eukaryote algal populations. The low-O2 CBB gene (cbbM) was most abundant near the lower oxycline boundary in the larger size fraction, coincident with purple sulfur bacteria populations. The reverse citric acid cycle gene (aclB) was equally abundant in both size fractions in the deepest photic zone, coinciding with green sulfur bacteria populations. Methane coenzyme reductase A (mcrA), of anaerobic methane cyclers, was most abundant at the lower oxycline boundary in both size fractions, coinciding with Methanoregula populations. δ13 CPOC values overlapped with the high-O2 CBB fixation range except for two negative excursions near the lower oxycline boundary, likely reflecting assimilation of isotopically-depleted groundwater-derived carbon by autotrophs and sulfate-reducers. Throughout aphotic waters, δ13 CPOC values of the large size fraction became 13 C-enriched, likely reflecting abundant purple sulfur bacterial aggregates. Eukaryote algae- or cyanobacteria-like isotopic signatures corresponded with increases in cbbL, cbbM, and aclB, and enrichment of exopolymer-rich prokaryotic photoautotrophs aggregates. Results suggest that δ13 CPOC values of preserved sediments from areas of the Proterozoic Ocean with sulfidic photic zones may reflect a mixture of alternate carbon-fixing populations exported from the deep photic zone, challenging the paradigm that sedimentary stable carbon isotope values predominantly reflect oxygenic photosynthesis from surface waters.

The photic-aphotic divide is a strong ecological and evolutionary force determining the distribution of ciliates (Alveolata, Ciliophora) in the ocean.

The bulk of knowledge on marine ciliates is from shallow and/or sunlit waters. We studied ciliate diversity and distribution across epi- and mesopelagic oceanic waters, using DNA metabarcoding and phylogeny-based metrics. We analyzed sequences of the 18S rRNA gene (V4 region) from 369 samples collected at 12 depths (0-1,000 m) at the Bermuda Atlantic Time-series Study site of the Sargasso Sea (North Atlantic) monthly for 3 years. The comprehensive depth and temporal resolutions analyzed led to three main findings. First, there was a gradual but significant decrease in alpha-diversity (based on Faith's phylogenetic diversity index) from surface to 1,000-m waters. Second, multivariate analyses of beta-diversity (based on UniFrac distances) indicate that ciliate assemblages change significantly from photic to aphotic waters, with a switch from Oligotrichea to Oligohymenophorea prevalence. Third, phylogenetic placement of sequence variants and clade-level correlations (EPA-ng and GAPPA algorithms) show Oligotrichea, Litostomatea, Prostomatea, and Phyllopharyngea as anti-correlated with depth, while Oligohymenophorea (especially Apostomatia) have a direct relationship with depth. Two enigmatic environmental clades include either prevalent variants widely distributed in aphotic layers (the Oligohymenophorea OLIGO5) or subclades differentially distributed in photic versus aphotic waters (the Discotrichidae NASSO1). These results settle contradictory relationships between ciliate alpha-diversity and depth reported before, suggest functional changes in ciliate assemblages from photic to aphotic waters (with prevalence of algivory and mixotrophy versus omnivory and parasitism, respectively), and indicate that contemporary taxon distributions in the vertical profile have been strongly influenced by evolutionary processes. Integration of DNA sequences with organismal data (microscopy, functional experiments) and development of databases that link these sources of information remain as major tasks to better understand ciliate diversity, ecological roles, and evolution in the ocean.

Astronomically controlled deep-sea life in the Late Cretaceous reconstructed from ultra-high-resolution inoceramid shell archives.

The periodicity of the mutual position of celestial bodies in the Earth-Moon-Sun system is crucial to the functioning of life on Earth. Biological rhythms affect most of the processes inside organisms, and some can be recorded in skeletal remains, allowing one to reconstruct the cycles that occur in nature deep in time. In the present study, we have used ultra-high-resolution elemental ratio scans of Mg/Ca, Sr/Ca and Mn/Ca from the fossil, ca. 70 Ma old inoceramid bivalve Inoceramus (Platyceramus) salisburgensis from deep aphotic water and identified a clear regularity of repetition of the geochemical signal every of ~0.006 mm. We estimate that the shell accretion rate is on average ~0.4cm of shell thickness per lunar year. Visible light-dark lamination, interpreted as a seasonal signal corresponding to the semilunar-related cycle, gives a rough shell age estimate and growth rate for this large bivalve species supported by a dual feeding strategy. We recognize a biological clock that follows either a semilunar (model A) or a tidal (model B) cycle. This cycle of tidal dominance seems to fit better considering the biological behaviour of I. (P.) salisburgensis, including the estimated age and growth rate of the studied specimens. We interpret that the major control in such deep-sea environment, well below the photic zone and storm wave base, was due to barotropic tidal forces, thus changing the water pressure.

Pulsed export of carbon in the north-western Mediterranean Sea

The short term (hourly scale) variability of heterotrophic prokaryote (HP) vertical distribution and respiratory activity, was investigated in the north-western (NW) Mediterranean Sea. HP vertical distribution was determined on board by flow cytometry analysis of seawater samples collected by series of CTD casts. Cell counts and viability were determined for all samples. HP respiratory rates were determined later in the laboratory from filtered seawater samples (23 dm3) from 300–1 150-m depth. The average cell viability was 94.8%±2.2% (n=240). There was no accumulation of dead cells, due to quick decay of damaged cells. In the epipelagic layer, three HP groups were distinguished, two (HNA1, HNA2) whose cells exhibited a high nucleic acid content and one (LNA) with low nucleic acid content cells. HNA2 was most populated at 50 m but not detected at 90 m and below, presumably aerobic anoxygenic photoheterotrophic bacteria (AAPs). The variability in HP abundance was mainly confined in the upper 80 m. A few secondary peaks of HP abundance were observed (80–150 m) in connection with abundance troughs in the surface layer. HP cells were continuously present in a wide layer around 500 m (mean 191×103 cells/cm3). Below this layer, HP abundance randomly exhibited peaks, coupled to respiratory rate peaks. The HP abundance and variability in the water column was suppressed during a strong wind event. The observed sporadic variability was tentatively interpreted through a pulsed carbon-export mechanism induced by the microorganism production of dissolved polysaccharides, followed by flocculation and rapid sinking. This mechanism would thus contribute to (i) preventing organic matter accumulation in the epipelagic layer, (ii) seeding the water column with live HP cells, and (iii) supplying the aphotic water column with fresh and labile organic matter. This important vertical flux mechanism needs further observations and modelling.

Deciphering the composite morphological diversity ofLophelia pertusa, a cosmopolitan deep‐water ecosystem engineer

AbstractCold‐water coral reefs constitute important biodiversity hotspots in aphotic waters around the world. The complex, highly variable morphology of the reef habitat‐forming species has important implications for the communities they harbor and for the physical processes occurring therein.Lophelia pertusa(Desmophyllum pertusum) is one of the most common reef‐building cold‐water corals, but its morphological diversity has never been characterized on a broad scale. We qualitatively and quantitatively explored the patterns of morphological variation of this species over a wide geographic and ecological range, addressing corallite and colony traits and their interrelation. Geographic variation is evident at both corallite and colony level, although with distinct trends. By linking branching patterns to colony morphology, we identified three main morphotypes (asymmetrical, bushy, and columnar) with substantial geometric and architectural differences, which suggest high functional diversity of cold‐water coral reefs across regions. Colony morphology appears strongly governed by asexual budding of individual polyps, but largely decoupled from corallite morphology. We hypothesize that colony morphology is primarily driven by local hydrodynamic conditions and associated food supply.

Changes in microbial community phylogeny and metabolic activity along the water column uncouple at near sediment aphotic layers in fjords

Fjords are semi-enclosed marine systems with unique physical conditions that influence microbial community composition and structure. Pronounced organic matter and physical condition gradients within fjords provide a natural laboratory for the study of changes in microbial community structure and metabolic potential in response to environmental conditions. Photosynthetic production in euphotic zones sustains deeper aphotic microbial activity via organic matter sinking, augmented by large terrestrial inputs. Previous studies do not consider both prokaryotic and eukaryotic communities when linking metabolic potential and activity, community composition, and environmental gradients. To address this gap we profiled microbial functional potential (Biolog Ecoplates), bacterial abundance, heterotrophic production (3H-Leucine incorporation), and prokaryotic/eukaryotic community composition (16S and 18S rRNA amplicon gene sequencing). Similar factors shaped metabolic potential, activity and community (prokaryotic and eukaryotic) composition across surface/near surface sites. However, increased metabolic diversity at near bottom (aphotic) sites reflected an organic matter influence from sediments. Photosynthetically produced particulate organic matter shaped the upper water column community composition and metabolic potential. In contrast, microbial activity at deeper aphotic waters were strongly influenced by other organic matter input than sinking marine snow (e.g. sediment resuspension of benthic organic matter, remineralisation of terrestrially derived organic matter, etc.), severing the link between community structure and metabolic potential. Taken together, different organic matter sources shape microbial activity, but not community composition, in New Zealand fjords.

'Ten Years After'-a long-term settlement and bioerosion experiment in an Arctic rhodolith bed (Mosselbukta, Svalbard).

Rhodolith beds and bioherms formed by ecosystem engineering crustose coralline algae support the northernmost centres of carbonate production, referred to as polar cold-water carbonate factories. Yet, little is known about biodiversity and recruitment of these hard-bottom communities or the bioeroders degrading them, and there is a demand for carbonate budgets to include respective rates of polar carbonate build-up and bioerosion. To address these issues, a 10-year settlement and bioerosion experiment was carried out at the Arctic Svalbard archipelago in and downslope of a rhodolith bed. The calcifiers recorded on experimental settlement tiles (56 taxa) were dominated by bryozoans, serpulids and foraminiferans. The majority of the bioerosion traces (30 ichnotaxa) were microborings, followed by attachment etchings and grazing traces. Biodiversity metrics show that calcifier diversity and bioerosion ichnodiversity are both elevated in the rhodolith bed, if compared to adjacent aphotic waters, but these differences are statistically insignificant. Accordingly, there were only low to moderate dissimilarities in the calcifier community structure and bioerosion trace assemblages between the two depth stations (46 and 127m), substrate orientations (up- and down-facing) and substrate types (PVC and limestone), in that order of relevance. In contrast, surface coverage as well as the carbonate accretion and bioerosion rates were all significantly elevated in the rhodolith bed, reflecting higher abundance or size of calcifiers and bioerosion traces. All three measures were highest for up-facing substrates at 46m, with a mean coverage of 78.2% (on PVC substrates), a mean accretion rate of 24.6gm-2 year-1 (PVC), and a mean bioerosion rate of -35.1gm-2 year-1 (limestone). Differences in these metrics depend on the same order of factors than the community structure. Considering all limestone substrates of the two platforms, carbonate accretion and bioerosion were nearly in balance at a net rate of -2.5gm-2 year-1 . A latitudinal comparison with previous settlement studies in the North Atlantic suggests that despite the harsh polar environment there is neither a depletion in the diversity of hard-bottom calcifier communities nor in the ichnodiversity of grazing traces, attachment etchings and microborings formed by organotrophs. In contrast, microborings produced by phototrophs are strongly depleted because of limitations in the availability of light (condensed photic zonation, polar night, shading by sea ice). Also, macroborings were almost absent, surprisingly. With respect to carbonate production, the Svalbard carbonate factory marks the low end of a latitudinal gradient while bioerosion rates are similar or even higher than at comparable depth or photic regime at lower latitudes, although this might not apply to shallow euphotic waters (not covered in our experiment), given the observed depletion in bioeroding microphytes and macroborers. While echinoid grazing is particularly relevant for the bioerosion in the rhodolith bed, respective rates are far lower than those reported from tropical shallow-water coral reefs. The slow pace of carbonate production but relatively high rates of bioerosion (both promoted by low carbonate supersaturation states in Arctic waters), in concert with high retention of skeletal carbonates on the seafloor and no calcite cements forming in open pore space created by microborers, suggest a low fossilisation potential for polar carbonates, such as those formed in the Mosselbukta rhodolith beds.

Dark Reduction Drives Evasion of Mercury From the Ocean

Much of the surface water of the ocean is supersaturated in elemental mercury (Hg0) with respect to the atmosphere, leading to sea-to-air transfer or evasion. This flux is large, and nearly balances inputs from the atmosphere, rivers and hydrothermal vents. While the photochemical production of Hg0 from ionic and methylated mercury is reasonably well-studied and can produce Hg0 at fairly high rates, there is also abundant Hg0 in aphotic waters, indicating that other important formation pathways exist. Here, we present results of gross reduction rate measurements, depth profiles and diel cycling studies to argue that dark reduction of Hg2+ is also capable of sustaining Hg0 concentrations in the open ocean mixed layer. In locations where vertical mixing is deep enough relative to the vertical penetration of UV-B and photosynthetically active radiation (the principal forms of light involved in abiotic and biotic Hg photoreduction), dark reduction will contribute the majority of Hg0 produced in the surface ocean mixed layer. Our measurements and modeling suggest that these conditions are met nearly everywhere except at high latitudes during local summer. Furthermore, the residence time of Hg0 in the mixed layer with respect to evasion is longer than that of redox, a situation that allows dark reduction-oxidation to effectively set the steady-state ratio of Hg0 to Hg2+ in surface waters. The nature of these dark redox reactions in the ocean was not resolved by this study, but our experiments suggest a likely mechanism or mechanisms involving enzymes and/or important redox agents such as reactive oxygen species and manganese (III).

Cold-water coral reefs thriving under hypoxia

Reefs formed by scleractinian cold-water corals represent unique biodiversity hot spots in the deep sea, preferring aphotic water depths of 200–1000 m. The distribution of the most prominent reef-building species Lophelia pertusa is controlled by various environmental factors including dissolved oxygen concentrations and temperature. Consequently, the expected ocean deoxygenation and warming triggered by human-induced global change are considered as a serious threat to cold-water coral reefs. Here, we present results on recently discovered reefs in the SE Atlantic, where L. pertusa thrives in hypoxic and rather warm waters. This sheds new light on its capability to adapt to extreme conditions, which is facilitated by high surface ocean productivity, resulting in extensive food supply. Putting our data in an Atlantic-wide perspective clearly demonstrates L. pertusa’s ability to develop population-specific adaptations, which are up to now hardly considered in assessing its present and future distributions.

Direct Detection of Heterotrophic Diazotrophs Associated with Planktonic Aggregates

N2 fixation by planktonic heterotrophic diazotrophs is more wide spread than previously thought, including environments considered “unfavorable” for diazotrophy. These environments include a substantial fraction of the aquatic biosphere such as eutrophic estuaries with high ambient nitrogen concentrations and oxidized aphotic water. Different studies suggested that heterotrophic diazotrophs associated with aggregates may promote N2 fixation in such environments. However, this association was never validated directly and relies mainly on indirect relationships and different statistical approaches. Here, we identified, for the first time, a direct link between active heterotrophic diazotrophs and aggregates that comprise polysaccharides. Our new staining method combines fluorescent tagging of active diazotrophs by nitrogenase-immunolabeling, polysaccharides staining by Alcian blue or concanavalin-A, and total bacteria via nucleic-acid staining. Concomitant to N2 fixation rates and bacterial activity, this new method provided specific localization of heterotrophic diazotrophs on artificial and natural aggregates. We postulate that the insights gained by this new visualization approach will have a broad significance for future research on the aquatic nitrogen cycle, including environments in which diazotrophy has traditionally been overlooked.

Diversity and Coding Potential of the Microbiota in the Photic and Aphotic Zones of Tropical Man-Made Lake with Intensive Aquaculture Activities: a Case Study on Temengor Lake, Malaysia.

Although freshwater biomes cover less than 1% of the Earth's surface, they have disproportionate ecological significances. Attempts to study the taxonomy and function of freshwater microbiota are currently limited to samples collected from temperate lakes. In this study, we investigated samples from the photic and aphotic of an aquaculture site (disturbed) of Temengor Lake, a tropical lake in comparison with the undisturbed site of the lake using 16S rRNA amplicon and shotgun metagenomic approaches. Vertical changes in bacterial community composition and function of the Temengor Lake metagenomes were observed. The photic water layer of Temengor Lake was dominated by typical freshwater assemblages consisting of Proteobacteria, Actinobacteria, Bacteroidetes, Verrucomicrobia, and Cyanobacteria lineages. On the other hand, the aphotic water featured in addition to Proteobacteria, Bacteroidetes, Verrucomicrobia, and two more abundant bacterial phyla that are typically ubiquitous in anoxic habitats (Chloroflexi and Firmicutes). The aphotic zone of Temengor Lake exhibited genetic potential for nitrogen and sulfur metabolisms for which terminal electron acceptors other than oxygen are used in the reactions. The aphotic water of the disturbed site also showed an overrepresentation of genes associated with the metabolism of carbohydrates, likely driven by the enrichment of nutrient resulting from aquaculture activities at the site. The results presented in this study can serve as a basis for understanding the structure and functional capacity of the microbial communities in the photic and aphotic zones/water layers of tropical man-made lakes.

A phylogenomic and ecological analysis of the globally abundant Marine Group II archaea (Ca. Poseidoniales ord. nov.)

Marine Group II (MGII) archaea represent the most abundant planktonic archaeal group in ocean surface waters, but our understanding of the group has been limited by a lack of cultured representatives and few sequenced genomes. Here, we conducted a comparative phylogenomic analysis of 270 recently available MGII metagenome-assembled genomes (MAGs) to investigate their evolution and ecology. Based on a rank-normalised genome phylogeny, we propose that MGII is an order-level lineage for which we propose the name Candidatus Poseidoniales (after Gr. n. Poseidon, God of the sea), comprising the families Candidatus Poseidonaceae fam. nov. (formerly subgroup MGIIa) and Candidatus Thalassarchaeaceae fam. nov. (formerly subgroup MGIIb). Within these families, 21 genera could be resolved, many of which had distinct biogeographic ranges and inferred nutrient preferences. Phylogenetic analyses of key metabolic functions suggest that the ancestor of Ca. Poseidoniales was a surface water-dwelling photoheterotroph that evolved to occupy multiple related ecological niches based primarily on spectral tuning of proteorhodopsin genes. Interestingly, this adaptation appears to involve an overwrite mechanism whereby an existing single copy of the proteorhodopsin gene is replaced by a horizontally transferred copy, which in many instances should allow an abrupt change in light absorption capacity. Phototrophy was lost entirely from five Ca. Poseidoniales genera coinciding with their adaptation to deeper aphotic waters. We also report the first instances of nitrate reductase in two genera acquired via horizontal gene transfer (HGT), which was a potential adaptation to oxygen limitation. Additional metabolic traits differentiating families and genera include flagellar-based adhesion, transporters, and sugar, amino acid, and peptide degradation. Our results suggest that HGT has shaped the evolution of Ca. Poseidoniales to occupy a variety of ecological niches and to become the most successful archaeal lineage in ocean surface waters.

Bioluminescent-like squamation in the galeomorph shark Apristurus ampliceps (Chondrichthyes: Elasmobranchii)

Galeomorph sharks constitute the most taxonomically and ecologically diverse superorder of living selachians. Despite comprising several typically deep-water taxa, no bioluminescent species have been reported in this group so far. Interestingly, the study of shark squamation has been revealed in recent years to be a good proxy for inferring some ecological aspects of poorly known species. In particular, the high morphological specificity of the dermal denticles and the squamation patterns of all currently-known bioluminescent sharks could constitute a potential tool for predicting bioluminescence in both fossil and living taxa. Following this idea, we provide the first evidence supporting the possible existence of bioluminescence among galeomorph sharks by means of the quantitative study of Apristurus ampliceps squamation pattern. Classical morphometric analysis and multivariate statistical procedures have allowed us to determine that A. ampliceps squamation, composed mainly of bristle-shaped dermal denticles, is highly convergent with that of the bioluminescent shark Etmopterus spinax. The ecology of A. ampliceps, being a species that exclusively inhabits aphotic waters, is in agreement with such a morphofunctional interpretation, but finding photophores is imperative to confirm this prediction.

Winter mixing impacts gene expression in marine microbial populations in the Gulf of Aqaba

AME Aquatic Microbial Ecology Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsSpecials AME 80:223-242 (2017) - DOI: https://doi.org/10.3354/ame01854 Winter mixing impacts gene expression in marine microbial populations in the Gulf of Aqaba D. Miller1, U. Pfreundt2,3, S. Hou2, S. C. Lott2, W. R. Hess2, I. Berman-Frank1,* 1Mina and Everard Goodman Faculty of Life Sciences, Bar Ilan University, Ramat Gan, Israel 2Genetics and Experimental Bioinformatics, Faculty of Biology, University of Freiburg, Schänzlestr. 1, 79104 Freiburg, Germany 3Present address: ETH Zürich, Department of Civil, Environmental and Geomatic Engineering, Institute of Environmental Engineering, Stefano-Franscini-Platz 5, 8093 Zürich, Switzerland *Corresponding author: ilana.berman-frank@biu.ac.il ABSTRACT: In aquatic systems, changes in temperature and irradiance fundamentally characterize the water column and regulate microbial population structure and function. In systems with stable thermal stratification, the warm surface mixed layer is typically nutrient impoverished, limiting biological production. In periods of destratification, convective mixing of the water column exposes the microorganisms inhabiting these mixed systems to rapid variations in light availability and spectra. We explored the impact of winter deep-mixing (500 m deep mixed layer) on microbial communities from the surface (2.5 m) and the aphotic waters (440 m) in the Gulf of Aqaba by examining changes in both population composition and function via DNA and RNA sequencing. The greatest fraction of 16S sequences was assigned to Euryarchaeota, while metatranscriptomes were dominated by Synechococcus transcripts. Community composition was highly similar at both depths, yet transcription profiles differed. Phototrophic organisms found at the photic surface overexpressed genes related to catabolism and energy metabolism, while genes affiliated with biosynthesis were overexpressed at the aphotic depth. Similar transcriptional trends were observed in the non-photoautotrophs SAR11, Euryarchaeota, and Thaumarchaeota, with niche partitioning based on differential utilization of nitrogen and phosphorus occurring between the 2 archaeal groups. We did not detect upregulated expression of cyanobacterial genes indicative of mixotrophy or glycogen metabolism in the aphotic zone, suggesting they survive the aphotic period by utilizing photosynthates produced in the photic zone. Indications for a mixotrophic lifestyle were observed for prasinophytes, with genes related to phagocytosis overexpressed at the aphotic depth compared with the surface. KEY WORDS: Deep mixing · Red Sea · Metatranscriptomics · Cyanobacteria · SAR11 · Archaea · Prasinophytes Full text in pdf format Supplement 1Supplement 2 PreviousNextCite this article as: Miller D, Pfreundt U, Hou S, Lott SC, Hess WR, Berman-Frank I (2017) Winter mixing impacts gene expression in marine microbial populations in the Gulf of Aqaba. Aquat Microb Ecol 80:223-242. https://doi.org/10.3354/ame01854 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in AME Vol. 80, No. 3. Online publication date: November 24, 2017 Print ISSN: 0948-3055; Online ISSN: 1616-1564 Copyright © 2017 Inter-Research.

Basin‐wide N2 fixation in the deep waters of the Mediterranean Sea

AbstractRecent findings indicate that N2 fixation is significant in aphotic waters, presumably due to heterotrophic diazotrophs depending on organic matter for their nutrition. However, the relationship between organic matter and heterotrophic N2 fixation remains unknown. Here we explore N2 fixation in the deep chlorophyll maximum and underneath deep waters across the whole Mediterranean Sea and relate it to organic matter composition, characterized by optical and molecular methods. Our N2 fixation rates were in the range of those previously reported for the euphotic zone of the Mediterranean Sea (up to 0.43 nmol N L−1 d−1) and were significantly correlated to the presence of relatively labile organic matter with fluorescence and molecular formula properties representative for peptides and unsaturated aliphatics and associated with the presence of more oxygenated ventilated water masses. Finally, and despite that the aphotic N2 fixation contributes largely to total water column diazotrophic activity (&gt;50%), its contribution to overall nitrogen inputs to the basin is negligible (&lt;0.5%).

Contrasting distributions of dissolved gaseous mercury concentration and evasion in the North Pacific Subarctic Gyre and the Subarctic Front

The distribution of dissolved gaseous mercury (DGM) and the oxidation–reduction processes of mercury (Hg) in the surface and subsurface ocean are currently understudied despite their importance in ocean–atmosphere interactions. We investigated the Hg(0) evasion and the DGM distribution at water depths of 2–500m in the Subarctic Front, Western Subarctic Gyre, and Bering Sea of the Northwestern Pacific. The mean DGM concentration in the surface mixed water (<10m) and the mean Hg(0) evasion flux were significantly higher in the Subarctic Front (125±5.0fM and 15pmolm−2h−1, respectively), which typically has lower nutrient levels and higher primary production, than in the Western Subarctic Gyre and the Bering Sea (74±18fM and 3.2±1.2pmolm−2h−1, respectively). The variation in the chlorophyll-a concentration and extracellular protease activity predicted 54% and 48% of the DGM variation, respectively, in the euphotic zone (2–50m). The DGM concentration in aphotic intermediate water (415±286fM) was positively correlated to the apparent oxygen utilization (AOU; r2=0.94 and p<0.001 for the Western Subarctic Gyre and the Bering Sea; r2=0.61 and p=0.01 for the Subarctic Front), emphasizing the importance of microbial oxidation of organic matter. The DGM-to-AOU ratio in aphotic water was significantly (p<0.05, ANCOVA) higher at the Western Subarctic Gyre and Bering Sea sites (2.5±0.14) than the ratio at the Subarctic Front sites (0.89±0.27) that mainly consisted of newly formed North Pacific Intermediate Water. The overall results imply that variation of DGM and Hg(0) evasion is closely linked to primary production in euphotic water and organic remineralization in aphotic intermediate water. The oceanic alterations in these factors may induce significant modification in Hg redox speciation in the Northwestern Pacific.

Impact of eutrophication and climate change on fish and zoobenthos in coastal waters of the Baltic Sea

Coastal waters are important in linking zoobenthos and fish, as many coastal fishes feed on benthic prey in these habitats. Major drivers, such as eutrophication and climate change, may alter this link, whereas shifts in the importance of these drivers may induce different responses in zoobenthos and fish, respectively, potentially changing productivity of coastal ecosystems. The aim of this study was to assess in which way abundance and distribution of benthic-feeding fish and biomass of zoobenthos have changed over time in response to eutrophication (Secchi depth) and climate change (temperature and salinity), respectively. This was done by analyzing gross changes in the responses over three decades (1983–2012) and across depth zones. Eutrophication and climate change caused different though specific impacts on fish and zoobenthos, respectively. Throughout the 1980s, increasing benthic-feeding fish abundance in shallow waters (<6 m) was primarily attributed to eutrophication (decreasing Secchi depth), implying increased system productivity. During the 2000s, the effect of eutrophication levelled out, whereas temperature caused contrasting development of fish abundance at different depth zones. Shallow waters had lower fish abundances during warm years compared with colder ones, while the abundance increased in deeper, aphotic waters (6–20 m). In the deep waters, zoobenthos showed a contrasting, declining trend in biomass, coinciding with the decrease in salinity during the 2000s. This suggests altered ecosystem productivity and potential food shortage for benthic-feeding fish exploring the deep waters. An intensification of these trends is likely in the future, as climate change scenarios suggest further temperature increase and salinity decrease.

Archaea and Bacteria in deep lake hypolimnion: in situ dark inorganic carbon uptake

The interest for microorganisms inhabiting the hypolimnion and for their role in biogeochemical cycles of lakes is considerable, but knowledge is far from complete. The presence of chemolithoautotrophic Bacteria and mesophilic Archaea (e.g., Thaumarchaeota) assimilating inorganic carbon in the deep hypolimnion of lakes has been ascertained. We measured, for the first time at 350 m in Lake Maggiore (Northern Italy), the prokaryotic in situ dark [14C]HCO3 incorporation with a new custom-made apparatus, which takes samples and adds tracers in situ. Thereby stress factors affecting prokaryotes during sample recovery from the depth were avoided. We tested the new instrument at different depths and conditions, performing parallel conventional on board incubations. We found that dark [14C]HCO3 incorporations had lower standard deviation in in situ incubations with respect to the on board ones, but their means were not statistically different. At 350 m we estimated an uptake of 187.7±15 μg C m–3 d–1, which is in line with the published uptake rates in aquatic systems. By inhibiting the bacterial metabolism, we found that Archaea were responsible for 28% of the total CO2 uptake. At the same depth, Thaumarchaeota, on average, constituted 11% of total DAPI counts. Dark [14C]HCO3 incorporation integrated along the aphotic water column was 65.8±5.2 mg C m–2 d–1 which corresponds to 87% of picophytoplanktonic autotrophic fixation in the euphotic layer. This study provides the first evidence of Bacteria and Archaea dark CO2 fixation in the deep hypolimnion of a subalpine lake and indicates a potentially significant prokaryotic CO2 sink.

Aphotic N2 Fixation in the Eastern Tropical South Pacific Ocean

We examined rates of N2 fixation from the surface to 2000 m depth in the Eastern Tropical South Pacific (ETSP) during El Niño (2010) and La Niña (2011). Replicated vertical profiles performed under oxygen-free conditions show that N2 fixation takes place both in euphotic and aphotic waters, with rates reaching 155 to 509 µmol N m−2 d−1 in 2010 and 24±14 to 118±87 µmol N m−2 d−1 in 2011. In the aphotic layers, volumetric N2 fixation rates were relatively low (<1.00 nmol N L−1 d−1), but when integrated over the whole aphotic layer, they accounted for 87–90% of total rates (euphotic+aphotic) for the two cruises. Phylogenetic studies performed in microcosms experiments confirm the presence of diazotrophs in the deep waters of the Oxygen Minimum Zone (OMZ), which were comprised of non-cyanobacterial diazotrophs affiliated with nifH clusters 1K (predominantly comprised of α-proteobacteria), 1G (predominantly comprised of γ-proteobacteria), and 3 (sulfate reducing genera of the δ-proteobacteria and Clostridium spp., Vibrio spp.). Organic and inorganic nutrient addition bioassays revealed that amino acids significantly stimulated N2 fixation in the core of the OMZ at all stations tested and as did simple carbohydrates at stations located nearest the coast of Peru/Chile. The episodic supply of these substrates from upper layers are hypothesized to explain the observed variability of N2 fixation in the ETSP.