Anammox bacteria and the anaerobic oxidation of ammonium in the oxygen minimum zone off northern Chile

Anammox is the anaerobic oxidation of ammonium by nitrite or nitrate to yield N₂. This process, along with conventional denitrification, contributes to nitrogen loss in oxygen-deficient systems. Anammox is performed by a special group of bacteria belonging to the Planctomycetes phylum. However, information about the distribution, activity, and controlling factors of these anammox bacteria is still limited. Herein, we examine the phylogenetic diversity, vertical distribution, and activity of anammox bacteria in the coastal upwelling region and oxygen minimum zone off northern Chile. The phylogeny of anammox bacteria was studied using primers designed to specifically target 16S rRNA genes from Planctomycetes in samples taken during a cruise in 2004. Anammox bacteria-like sequences affiliated with Candidatus “Scalindua spp.” dominated the 16S rRNA gene clone library. However, 62% of the sequences subgrouped separately within this cluster and together with a single sequence retrieved from the suboxic zone of the freshwater Lake Tanganyika. The vertical distribution and activity of anammox bacteria were explored through CARD-FISH (fluorescence in situ hybridization with catalyzed reporter deposition) and ¹⁵N labeling incubations, respectively, at two different open-ocean stations during a second cruise in 2005. Anammox bacterial CARD-FISH counts (up to 3000 cells ml⁻¹) and activity (up to 5.75 nmol N₂ L⁻¹ d⁻¹) were only detected at the station subjected directly to the upwelling influence. Anammox cell abundance and activity were highest at 50 m depth, which is the upper part of the OMZ. In this layer, a high abundance of cyanobacteria and a marked nitrogen deficit were also observed. Thus, our results show the presence of a new subcluster within the marine anammox phylogeny and indicate high vertical variability in the abundance and activity of anammox bacteria that could be related to an intensification of carbon and nitrogen cycling in the upper part of the OMZ.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

Biomass size-spectra of macrobenthic communities in the oxygen minimum zone off Chile

Estimates of macrofaunal secondary production and normalized biomass size-spectra (NBSS) were constructed for macrobenthic communities associated with the oxygen minimum zone (OMZ) in four areas of the continental margin off Chile. The presence of low oxygen conditions in the Humboldt Current System (HCS) off Chile was shown to have important effects on the size structure and secondary production of the benthic communities living in this ecosystem. The distribution of normalized biomass by size was linear (log₂–log₂ scale) at all stations. The slope of the NBSS ranged from −0.481 to −0.908. There were significant differences between the slopes of the NBS-spectra from the stations located in the OMZ (slope = −0.837) and those located outside the OMZ (slope = −0.463) (p < 0.05). The results of this study suggest that low oxygen conditions (<0.5 ml L⁻¹) appear to influence biomass size-spectra, because small organisms are better able to satisfy their metabolic demands. The annual secondary production was higher off central Chile (6.8 g C m⁻² y⁻¹) than off northern Chile (2.02 g C m⁻² y⁻¹) and off southern Chile (0.83 g C m⁻² y⁻¹). A comparison with other studies suggests that secondary production in terms of carbon equivalents was higher than in other upwelling regions.     

Metazoan meiofauna within the oxygen-minimum zone off Chile: Results of the 2001-PUCK expedition

A quantitative study of metazoan meiofauna was carried out at continental shelf and slope stations affected by the oxygen-minimum zone in the eastern South Pacific off Chile. Densities of meiobenthos at the investigated stations off Antofagasta (22°S), Concepción (36°S), and Chiloé (42°S) ranged from 1282.1 to 8847.8 ind 10 cm⁻². Oxygen deficiency led only to average abundances, despite higher food availability and freshness at the corresponding sites. Sediment organic carbon, chlorophyll-a, and phaeopigment contents were used as measures of the input from water-column primary production, which accumulated at the oxygen-minimum zone stations. The highest abundances were found at a station with an oxygen content of 0.79 mL L⁻¹, which was slightly elevated from what is defined as oxygen minimum (0.5 mL L⁻¹). The most oxygenated site yielded the lowest densities. Meiofauna assemblages became more diverse with increasing bottom-water oxygenation, whereas nematodes were the most abundant taxon at every station, followed by annelids, copepods, and nauplii.     

Meiofaunal distributions on the Peru margin:: relationship to oxygen and organic matter availability

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12°S). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440–548 ind. 10 cm⁻² (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2–5 cm interval. At the deeper sites, about 50% were restricted to the top 1 cm. The importance of copepods and nauplii increased consistently with depth, reaching ∼12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02 ml l⁻¹ supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r=0.99, p<0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites.     

Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000

Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m⁻² d⁻¹ between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m⁻² d⁻¹ to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.     

Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean

Studies of nitrogen and phosphorus dynamics in the oligotrophic surface waters of the western North Atlantic Ocean have been constrained because ambient concentrations are typically at or below the detection limits of standard colorometric methods, except during periods of deep vertical mixing. Here we report the application of high-sensitivity analytical methods—determinations of nitrate plus nitrite (N+N) by chemiluminescence and soluble reactive phosphorus (SRP) by the magnesium induced co-precipitation (MAGIC) protocol—to surface waters along a transect from the Sargasso Sea at 26°N through the Gulf Stream at 37°N, including sampling at the JGOFS Bermuda Atlantic Time-series Study (BATS) station. The results were compared with data from the BATS program, and the HOT station in the Pacific Ocean, permitting cross-ecosystem comparisons. Microbial populations were analyzed along the transect, and an attempt was made to interpret their distributions in the context of the measured nutrient concentrations.Surface concentrations of N+N and SRP during the March 1998 transect separated into 3 distinct regions, with the boundaries corresponding roughly to the locations of the BATS station (∼31°N) and the Gulf Stream (∼37°N). Although N+N and SRP co-varied, the [N+N] : [SRP] molar ratios increased systematically from ∼1 to 10 in the southern segment, remained relatively constant at ∼40–50 between 31°N and 37°N, then decreased again systematically to ratios <10 north of the Gulf Stream. Dissolved organic N (DON) and P (DOP) dominated (⩾90%) the total dissolved N (TDN) and P (TDP) pools except in the northern portion of the transect. The [DON] : [DOP] molar ratios were relatively invariant (∼30–60) across the entire transect.Heterotrophic prokaryotes (operationally defined as “bacteria”), Prochlorococcus, Synechococcus, ultra- and nanophytoplankton, cryptophytes, and coccolithophores were enumerated by flow cytometry. The abundance of bacteria was well correlated with the concentration of SRP, and that of the ultra- and nanophytoplankton was well correlated with the concentration of N+N. The only group whose concentration was correlated with temperature was Prochlorococcus, and its abundance was unrelated to the concentrations of nutrients measured at the surface.We combined our transect results with time-series measurements from the BATS site and data from select depth profiles, and contrasted these North Atlantic data sets with time-series of N and P nutrient measurements from a station in the North Pacific subtropical gyre near Hawaii [Hawaii Ocean Time-series (HOT) site]. Two prominent differences are readily observed from this comparison. The [N+N] : [SRP] molar ratios are much less than 16 : 1 during stratified periods in surface waters at the BATS site, as is the case at the HOT site year round. However, following deep winter mixing, this ratio is much higher than 16 : 1 at BATS. Also, SRP concentrations in the upper 100 m at BATS fall in the range 1–10 nM during stratified periods, which is at least one order of magnitude lower than at the HOT site. That two ecosystems with comparable rates of primary and export production would differ so dramatically in their nutrient dynamics is intriguing, and highlights the need for detailed cross ecosystem comparisons.     

Nitrous oxide and N-nutrient cycling in the oxygen minimum zone off northern Chile

Measurements of dissolved gases (O₂, N₂O), nutrients (NO₃⁻, NO₂⁻, PO₄³⁻), and oceanographic variables were performed off northern Chile (∼21°S) between March 2000 and July 2004, in order to characterize the existing oxygen minimum zone (OMZ) and identify processes involved in N₂O cycling. Both N₂O and NO₃⁻ displayed sharp, shallow peaks with concentrations of up to 124 nM (1370% saturation) and 26 μM, respectively, in association with a strong oxycline that impinges on the euphotic zone. NO₂⁻ accumulation below the oxycline's base reached up to 9 μM. The vertical distribution of physical and chemical parameters and the existing relationships between apparent oxygen utilization (AOU), apparent N₂O production (ΔN₂O), and NO₃⁻ revealed three main layers within the upper OMZ. The first layer, or the upper part of the oxycline, is located between the base of the mixed layer and the mid-point of the oxycline (around σ_t=25.5 kg m⁻³). There the O₂ declines from ∼250 to ∼50 μM, and strong (but opposing) O₂ and NO₃⁻ gradients and their associated AOU–ΔN₂O and AOU–NO₃⁻ relationships indicate that nitrification produces N₂O and NO₃⁻ in the presence of light. The second layer, or lower part of the oxycline, represents the upper OMZ boundary and is located between the middle and the base of the oxycline (25.9<σ_t<26.1 kg m⁻³). In this layer NO₃⁻ reduction begins at O₂ levels ranging from ∼50 to ∼11 μM and accumulation of 41–68% of the ΔN₂O pool occurs. The accumulation of N₂O (but not of NO₂⁻ or NH₄⁺) and the observed AOU–ΔN₂O and AOU–NO₃⁻ relationships (which are opposite to those of the overlying first layer) suggest that a coupling between nitrification and NO₃⁻ reduction is involved in N₂O cycling in this second layer. The third layer is the OMZ core, where the O₂ concentration remains constant (O₂<11 μM). It coincides with σ_t>26.2 kg m⁻³, which is typical of Equatorial Subsurface Water (ESSW). In this layer, N₂O and NO₃⁻ continue to decrease, but a large NO₂⁻ accumulation is observed. Considering all the data, a biogeochemical model for the upper OMZ off northern of Chile is proposed, in which nitrification and denitrification differentially mediate N₂O cycling in each layer.     

Spatial and temporal variation in surfclam (Spisula solidissima) larval supply and settlement on the New Jersey inner shelf during summer upwelling and downwelling

To examine the relationship between near-bottom larval surfclam concentrations and surfclam settlement at an inner continental shelf site off New Jersey (USA), four consecutive sets of settlement experiments were carried out at three stations at the Long-term Ecosystem Observatory (LEO-15) from 14 to 31 July 1997 during upwelling and downwelling. Two inshore stations were on the landward and seaward sides of Beach Haven Ridge at ∼12 m depth, and a third station was 8 km further offshore at ∼20 m depth. In each experiment, four replicate trays of azoic sand from Beach Haven Ridge were placed flush with the seafloor and exposed for 3–7 days. Larval surfclam concentrations were measured every 4 h at 1 m above the bottom (mab) using Moored, Automated, Serial Zooplankton Pumps at the three stations. At all three stations, larval surfclam concentrations (1 mab) were low during upwelling, and higher during and after downwelling. Pulses of highest larval surfclam concentrations coincided with the initial arrival of downwelled warm water. In addition, larval surfclam concentrations were higher at the two inshore stations than at the offshore station. Larval surfclam settlement in the trays was higher during and following downwelling than during upwelling at one inshore station and at the offshore station. At the other inshore station (landward of Beach Haven Ridge), surfclam settlement did not increase during and following downwelling. Overall, surfclam settlement was higher inshore than offshore. The results indicate that spatial and temporal variation in larval surfclam supply was controlled by upwelling and downwelling circulation and that surfclam settlement was influenced by larval supply. Bottom flows across Beach Haven Ridge during a storm may have reduced larval surfclam settlement on the upcurrent side of the ridge, affecting initial densities on a small (∼1 km) scale.     

Distribution and burial of organic carbon in sediments from the Indian Ocean upwelling region off Java and Sumatra,Indonesia

Sediments were sampled and oxygen profiles of the water column were determined in the Indian Ocean off west and south Indonesia in order to obtain information on the production, transformation, and accumulation of organic matter (OM). The stable carbon isotope composition (δ¹³C_org) in combination with C/N ratios depicts the almost exclusively marine origin of sedimentary organic matter in the entire study area. Maximum concentrations of organic carbon (C_org) and nitrogen (N) of 3.0% and 0.31%, respectively, were observed in the northern Mentawai Basin and in the Savu and Lombok basins. Minimum δ¹⁵N values of 3.7‰ were measured in the northern Mentawai Basin, whereas they varied around 5.4‰ at stations outside this region. Minimum bottom water oxygen concentrations of 1.1 mL L^?1, corresponding to an oxygen saturation of 16.1%, indicate reduced ventilation of bottom water in the northern Mentawai Basin. This low bottom water oxygen reduces organic matter decomposition, which is demonstrated by the almost unaltered isotopic composition of nitrogen during early diagenesis. Maximum C_org accumulation rates (CARs) were measured in the Lombok (10.4 g C m^?2 yr^?1) and northern Mentawai basins (5.2 g C m^?2 yr^?1). Upwelling-induced high productivity is responsible for the high CAR off East Java, Lombok, and Savu Basins, while a better OM preservation caused by reduced ventilation contributes to the high CAR observed in the northern Mentawai Basin. The interplay between primary production, remineralisation, and organic carbon burial determines the regional heterogeneity. CAR in the Indian Ocean upwelling region off Indonesia is lower than in the Peru and Chile upwellings, but in the same order of magnitude as in the Arabian Sea, the Benguela, and Gulf of California upwellings, and corresponds to 0.1–7.1% of the global ocean carbon burial. This demonstrates the relevance of the Indian Ocean margin off Indonesia for the global OM burial.     

Depth-related distribution and abundance of seastars (Echinodermata: Asteroidea) in the Porcupine Seabight and Porcupine Abyssal Plain,N.E. Atlantic

The depth-related distribution of seastar (Echinodermata: Asteroidea) species between 150 and 4950 m in the Porcupine Seabight and Porcupine Abyssal Plain is described. 47 species of asteroid were identified from ∼14,000 individuals collected. The bathymetric range of each species is recorded. What are considered quantitative data, from an acoustically monitored epibenthic sledge and supplementary data from otter trawls, are used to display the relative abundance of individuals within their bathymetric range. Asteroid species are found to have very narrow centres of distribution in which they are abundant, despite much wider total adult depth ranges. Centres of distribution may be skewed. This might result from competition for resources or be related to the occurrence of favourable habitats at particular depths. The bathymetric distributions of the juveniles of some species extend outside the adult depth ranges. There is a distinct pattern of zonation with two major regions of faunal change and six distinct zones. An upper slope zone ranges from 150 to ∼700 m depth, an upper bathyal zone between 700 and 1100 m, a mid-bathyal zone from 1100 to1700 m and a lower bathyal zone between 1700 and 2500 m. Below 2500 m the lower continental slope and continental rise have a characteristic asteroid fauna. The abyssal zone starts at about 2800 m. Regions of major faunal change are identified at the boundaries of both upper and mid-bathyal zones and at the transition of bathyal to abyssal fauna. Diversity is greatest at ∼1800 m, decreasing with depth to ∼2600 m before increasing again to high levels at ∼4700 m.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Abundance of small individuals influences the effectiveness of processing techniques for deep-sea nematodes

Nematodes are the most abundant metazoans of deep-sea benthic communities, but knowledge of their distribution is limited relative to larger organisms. Whilst some aspects of nematode processing techniques, such as extraction, have been extensively studied, other key elements have attracted little attention. We compared the effect of (1) mesh size (63, 45, and 32 μm) on estimates of nematode abundance, biomass, and body size, and (2) microscope magnification (50× and 100×) on estimates of nematode abundance at bathyal sites (250–3100 m water depth) on the Challenger Plateau and Chatham Rise, south-west Pacific Ocean. Variation in the effectiveness of these techniques was assessed in relation to nematode body size and environmental parameters (water depth, sediment organic matter content, %silt/clay, and chloroplastic pigments). The 63-μm mesh retained a relatively low proportion of total nematode abundance (mean±SD=55±9%), but most of nematode biomass (90±4%). The proportion of nematode abundance retained on the 45-μm mesh in surface (0–1 cm) and subsurface (1–5 cm) sediment was significantly correlated (P<0.01) with %silt/clay (R²=0.39) and chloroplastic pigments (R²=0.29), respectively. Variation in median nematode body weight showed similar trends, but relationships between mean nematode body weight and environmental parameters were either relatively weak (subsurface sediment) or not significant (surface sediment). Using a low magnification led to significantly lower (on average by 43%) nematode abundance estimates relative to high magnification (P<0.001), and the magnitude of this difference was significantly correlated (P<0.05) with total nematode abundance (R²_p=0.53) and the number of small (≤250 μm length) individuals (R²_p=0.05). Our results suggest that organic matter input and sediment characteristics influence the abundance of small nematodes in bathyal communities. The abundance of small individuals can, in turn, influence abundance estimates obtained using different mesh sizes and microscope magnifications.     

Mesoscale distribution and advection of overwintering Calanus finmarchicus off the shelf of northern Norway

Data collected on a cruise in January 2008, using a laser optical plankton counter, conductivity–temperature–depth sensors, and a lowered acoustic Doppler current profiler, was used to study the mesoscale distribution and advection of overwintering Calanus finmarchicus in its deep water winter habitat off the shelf of northern Norway. The overwintering animals were generally located immediately below the Atlantic Water (AW) in Arctic Intermediate Water (AIW), in the 600–1200 m depth range. The depth of the interface between AW and AIW varied considerably in the area and this was clearly reflected in the C. finmarchicus distribution. Maximum abundance varied from about 80 ind m^?3 to more than 200 ind m^?3 at the different stations. Current measurements showed the richness of mesoscazle eddies, with speeds exceeding 70 cm s^?1 at the surface and rapidly decreasing with depth. In the main overwintering layer the eddy features were also clearly seen, but with speeds generally below 20 cm s^?1. C. finmarchicus were found in the whole survey area, but they were not homogeneously distributed. Advection of the copepods resulted in relatively high local rates of change in overwintering C. finmarchicus abundance with mean value of 8% per day in the area. It is clear that mesoscale physical processes greatly contribute to the variability in the abundance of overwintering C. finmarchicus off the shelf of northern Norway. The collected data are also a valuable addition to the generally sparse datasets on the C. finmarchicus winter distribution and the role of the Lofoten basin in the large scale system is also discussed.     

Iron fertilization of Trichodesmium off the west coast of Barbados: A one-dimensional numerical model

A one-dimensional quasi-Eulerian model, describing the biological and chemical interactions of autotrophic and heterotrophic plankton populations within the upper 100 m of the water column, was used to explain the relative impact of iron fertilization from Saharan dust on phosphorus cycling by the cyanobacterium, Trichodesmium spp. We examined the Caribbean surface waters off the west coast of Barbados using seasonal cases of dust delivery (summer peak) in relation to periods of elevated phosphorus stocks found in episodic pulses of Amazon River water advected past Barbados by the Guiana Current. The additional iron and phosphorus supplied by fluvial sources, available after biological depletion of nitrogen, could alleviate the growth limitation of Trichodesmium far downstream of the river plumes. The model results were compared with measurements made of Trichodesmium stocks 8 km off the west coast of Barbados. Modeled concentrations within the low-salinity Amazon plumes reached ∼7.4 mg chl m⁻², as seen at Barbados. In the absence of the low-salinity signal, the Trichodesmium biomass shifted to phosphorus limitation with little response to iron fertilization. This indicates that Trichodesmium stocks may indeed be mainly a function of phosphorus availability within the upper water column. Therefore, the supply of phosphorus from subtropical/tropical rivers, such as the Amazon and Orinoco, limits the potential cyanophyte response, despite adequate atmospheric iron delivery. This alters our view not only of the western North Atlantic nitrogen budget, but potentially several regions downstream of low N:P river systems.     

Vertical distribution of prokaryote production and abundance in the mesopelagic and bathypelagic layers of the Canada Basin,western Arctic: Implications for the mode and extent of organic carbon delivery

The vertical distributions of prokaryote heterotrophic production (³H-leucine incorporation rate) and abundance were investigated in the meso- and bathy-pelagic layers of the Canada Basin, western Arctic Ocean, during September 2009. Prokaryote production and abundance were high in the Pacific-origin water mass located in the upper mesopelagic layer (depth, 100–200 m). Below the halocline layer (depth, 300–3000 m), both the production and abundance decreased with depth, with log–log regression slopes of −1.33 and −0.77, respectively. Depth-integrated production and biomass in the meso- and bathy-pelagic layers was three- to five-fold lower than the corresponding values reported in the subpolar regions, whereas they were close to or lower than the corresponding values in oligotrophic subtropical regions. Prokaryote turnover times were estimated to be 1.1 and 6.1 years for meso- and bathy-pelagic layers, respectively, with the latter being among the longest turnover times reported for oceanic basins. We estimated prokaryote carbon demand in the water column (100–3000 m) to be on the order of 11 mg C m⁻² d⁻¹, which largely exceeds (by 38-fold) the sinking particulate organic carbon flux at depths of 120–200 m reported in the literature. This large carbon imbalance may be partly explained by organic carbon delivery by lateral intrusion of the Pacific-origin water mass into the upper mesopelagic layer.     

Simulated sequestration of anthropogenic carbon dioxide at a deep-sea site: Effects on nematode abundance and biovolume

One proposal for ameliorating global warming is to sequester large amounts of carbon dioxide in the deep ocean, but the environmental consequences of sequestration for sediment-dwelling animals are poorly known. In a previous publication, we reported that ∼80% of benthic copepods were killed in an experimental release of CO₂ off northern California at 3262 m. The effects of this release on nematodes are reported here. We examined samples of nematodes taken inside two ‘corrals’ into which CO₂ was directly injected (providing an extreme endpoint for CO₂ exposure) and taken near to and far from this CO₂ source. After 30 days, pore-water pH was unchanged (∼7.8) at the sediment–water interface far (∼40 m) from corrals, but pH profiles were reduced by ∼0.75 near (∼2 m) corrals. Corral pH was highly acidic (5.4 in a measurement from a subsequent experiment). Fifty randomly selected nematodes from each of four vertical layers from the 14 cores were photographed. They were assigned to a tail group (based on morphology), and individual biovolume was estimated from measurements of body length and width. Although nematode abundance (expressed as total nematodes and by tail group) was not affected, length, width, and individual biovolume significantly differed between near and far samples. Median nematode biovolume examined across tail group and core layer increased by ∼48% inside and near corrals. Differences between near and corral samples were always less than differences between near and far samples. However, nematode length:width ratio did not differ between near and far, and the shapes of length, width, and biovolume frequency distributions were similar in all samples. We postulate that the nematode community throughout the upper 3 cm suffered a high rate of mortality after exposure to CO₂, and that nematodes were larger because postmortem expansions in body length and width occurred. Decomposition rates were probably low and corpses did not disintegrate in 30 days. The observable effects of a reduction in pH to about 7.0 after 30 days were as great as an extreme pH reduction (5.4), suggesting that ‘moderate’ CO₂ exposure, compared to the range of exposures possible following CO₂ release, causes high mortality rates in the two most abundant sediment-dwelling metazoans (nematodes and copepods).     

Distribution and diversity of open-ocean,near-bottom macroplankton in the western Mediterranean: Analysis at different spatio-temporal scales

We analyzed the distribution, diversity, and composition of western Mediterranean macroplankton (excluding gelatinous taxa) in the water column over depths of ca. 550–850 m, with special attention to near-bottom (0–1.5 and ca. 5–77 m above the bottom, mab) levels, and including data from three areas (off the coasts of Catalonia, and to the NW, and SE of Mallorca, Balearic Islands) in the period 1991–2008. Spatio-temporal changes in macroplankton abundance were evaluated as follows: (i) by seasonal sampling in 2007 off the Catalonian coast, (ii) by comparing Catalonian and Balearic Island slopes, and (iii) by comparing a fixed station on the Catalonian slope (at 550–800 m depth) at decadal (1991/1992–2007/2008) time scales. Diversity (in terms of species richness, S) was greater (i) at ca. 5–77 mab than at 0–1.5 mab, (ii) over the insular slopes of the Balearic Island (around Mallorca) than over the mainland Catalonian slopes, and (iii) in the period 1991/1992 than in 2007, likely related to higher values of the North Atlantic Oscillation (NAO) index in 1991/1992. In most analyses species composition was strongly influenced by the degree of stratification and homogenization of the water column in summer–autumn and winter–spring respectively and by location (longitude). Changes consisted mainly of higher density of macroplankton (e.g. abundance of the dominant euphausiids Nematoscelis megalops, Meganyctiphanes norvegica and Euphausia krohni and of the fish Cyclothone braueri) between June and October, parallel to an increase in the T and S close to the bottom. This coincided with changes in the flow of Levantine intermediate water (LIW) in the area. Aggregation of adult forms of the dominant species close to the bottom in summer–autumn could be favored because summer is the period of highest density of food – copepods, mainly Calanus helgolandicus – near the bottom off the Catalan slope. The formation of a thermocline in the water column and the reinforcement of the permanent thermohaline front at the shelf-slope break during summer at ca. 400 m in the Balearic Basin may also enhance this tendency toward greater aggregation of deep macroplankton under stratified water column conditions.     

Depth zonation and bathymetric trends of deep-sea megafaunal scavengers of the Hawaiian Islands

The deep sea has been shown to exhibit strong depth zonation in species composition and abundance. Examination of these patterns can offer ecological insight into how organisms adapt and respond to changing environmental parameters that co-occur with depth. Here we provide the first tropical study on bathymetric zonation and other depth-related trends (size, abundance, and species richness) spanning shelf to abyssal depths of scavenging megafauna. Baited time-lapse free-vehicle cameras were used to examine the deep-sea benthic and demersal scavenging communities of the Hawaiian Islands, an area for which the biology and ecology have remained poorly studied below 2000 m. Twenty-two deployments ranging in depth from 250 to 4783 m yielded 37 taxa attracted to bait, including the first known occurrence of the family Zoarcidae in the Hawaiian Islands. Cluster analysis of Bray–Curtis similarity of species peak abundance (n_max) revealed four main faunal zones (250–500, 1000, 1500–3000, and ?4000 m) with significant separation (ANOSIM, global R=0.907, p=0.001) between designated depth groups. A major faunal break was identified at the 500–1000 m transition where species turnover was greatest, coinciding with the location of the local oxygen minimum zone. Dominance in species assemblage shifted from decapod crustaceans to teleosts moving from shallow to deeper faunal zones. Significant size differences in total length with depth were found for two of the four fish species examined. A logarithmic decline was observed in scavenger relative abundance with depth. Evidence of interaction between scavenging species was also noted between Synaphobranchus affinis and Neolithodes sp. (competition) and Histiobranchus sp. and aristeid shrimp (predation), suggesting that interactions between scavengers could influence indices of abundance generated from baited camera data.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.