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Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg C·m−2·d−1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional ∼225 mg C·m−2·d−1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.

Glacial meltwater from the western Antarctic Ice Sheet is hypothesized to be an important source of cryospheric iron, fertilizing the Southern Ocean, yet its trace-metal composition and factors that control its dispersal remain poorly constrained. Here we characterize meltwater iron sources in a heavily glaciated western Antarctic Peninsula (WAP) fjord. Using dissolved and particulate ratios of manganese to iron in meltwaters, porewaters, and seawater, we show that surface glacial melt and subglacial plumes contribute to the seasonal cycle of iron and manganese within a fjord still relatively unaffected by climate-change-induced glacial retreat. Organic ligands derived from the phytoplankton bloom and the glaciers bind dissolved iron and facilitate the solubilization of particulate iron downstream. Using a numerical model, we show that buoyant plumes generated by outflow from the subglacial hydrologic system, enriched in labile particulate trace metals derived from a chemically modified crustal source, can supply iron to the fjord euphotic zone through vertical mixing. We also show that prolonged katabatic wind events enhance export of meltwater out of the fjord. Thus, we identify an important atmosphere–ice–ocean coupling intimately tied to coastal iron biogeochemistry and primary productivity along the WAP.

Coastal upwelling regions are among the most productive marine ecosystems but may be threatened by amplified ocean acidification. Increased acidification is hypothesized to reduce iron bioavailability for phytoplankton thereby expanding iron limitation and impacting primary production. Here we show from community to molecular levels that phytoplankton in an upwelling region respond to short-term acidification exposure with iron uptake pathways and strategies that reduce cellular iron demand. A combined physiological and multi-omics approach was applied to trace metal clean incubations that introduced 1200 ppm CO 2 for up to four days . Although variable, molecular-level responses indicate a prioritization of iron uptake pathways that are less hindered by acidification and reductions in iron utilization. Growth, nutrient uptake, and community compositions remained largely unaffected suggesting that these mechanisms may confer short-term resistance to acidification; however, we speculate that cellular iron demand is only temporarily satisfied, and longer-term acidification exposure without increased iron inputs may result in increased iron stress. Coastal upwelling regions are among the most productive marine ecosystems but may be threatened by amplified ocean acidification. Here the authors show from community to molecular levels that phytoplankton in an upwelling region respond to short-term acidification exposure with iron uptake pathways and strategies that reduce cellular iron demand.

The distribution of dissolved iron in the southern California Current System (sCCS) is presented from seven research cruises between 2002 and 2006. Dissolved iron concentrations were generally low in most of the study area (<0.5 nM), although high mixed layer and water column dissolved iron concentrations (up to 8 nM) were found to be associated with coastal upwelling, both along the continental margin and some island platforms. A significant supply of iron was probably not from a deep remineralized source but rather from the continental shelf and bottom boundary layer as identified in previous studies along the central and northern California coast. With distance offshore, dissolved iron decreased more rapidly relative to nitrate in a transition zone 10–250 km offshore during spring and summer, resulting in relatively high ratios of nitrate:dissolved iron. Higher nitrate:dissolved iron ratios could be the result of utilization and scavenging in addition to an overall lower supply of iron relative to nitrate in the offshore transition zones. The low supply of iron leads to phytoplankton iron limitation and a depletion in silicic acid relative to nitrate in the coastal upwelling and transition zones of the sCCS.

Organic dissolved iron (dFe)–binding ligands were measured by competitive ligand exchange–adsorptive cathodic stripping voltammetry (CLE-ACSV) at multiple analytical windows (side reaction coefficient of salicylaldoxime, αFe(SA)₂ = 30, 60, and 100) in surface and benthic boundary layer (BBL) samples along the central California coast during spring and summer. The weakest ligands were detected in the BBL at the lowest analytical window with average log K FeL, Fe ' cond = 10.2 ± 0.4 in the summer and 10.8 ± 0.2 in the spring. Between 3% and 18% of the dFe complexation in the BBL was accounted for by HS, which were measured separately in samples by ACSV and may indicate a source of dFe-binding ligands from San Francisco Bay. The strongest ligands were found in nearshore spring surface waters at the highest analytical window with average log K Fel, Fe ' cond = 11.9 ± 0.3 , and the concentrations of these ligands declined rapidly offshore. The ligand pools in the surface and BBL waters were distinct from each other based on principal components analysis, with variances in the BBL ligand pool explained by sample location, and variance in surface waters explained by water mass. The use of multiple analytical window analysis elucidated several distinct iron-binding ligand pools, each with unique distributions in the central California Current system.

Organic ligands form strong complexes with many trace elements in seawater. Various metals can compete for the same ligand chelation sites, and the final speciation of bound metals is determined by relative binding affinities, concentrations of binding sites, uncomplexed metal concentrations, and association/dissociation kinetics. Different ligands have a wide range of metal affinities and specificities. However, the chemical composition of these ligands in the marine environment remains poorly constrained, which has hindered progress in modeling marine metal speciation. In this study, we detected and characterized natural ligands that bind copper (Cu) and nickel (Ni) in the eastern South Pacific Ocean with liquid chromatography tandem inductively coupled plasma mass spectrometry (LC-ICPMS), and high resolution electrospray ionization mass spectrometry (ESIMS). Dissolved Cu, Ni, and ligand concentrations were highest near the coast. Chromatographically unresolved polar compounds dominated ligands isolated near the coast by solid phase extraction. Offshore, metal and ligand concentrations decreased, but several new ligands appeared. One major ligand was detected that bound both Cu2+ and Ni2+. Based on accurate mass and fragmentation measurements, this compound has a molecular formula of [C20H21N4O8S2 + M]+ (M = metal isotope) and contains several azole-like metal binding groups. Additional lipophilic Ni complexes were also present only in oligotrophic waters, with masses of 649, 698, and 712 m/z (corresponding to the 58Ni metal complex). Molecular formulae of [C32H54N3O6S2Ni]+ and [C33H56N3O6S2Ni]+ were determined for two of these compounds. Addition of Cu and Ni to the samples also revealed the presence of additional compounds that can bind both Ni and Cu. Although these specific compounds represent a small fraction of the total dissolved Cu and Ni pool, they highlight the compositional diversity and spatial heterogeneity of marine Ni and Cu ligands, as well as variability in the extent to which different metals in the same environment compete for ligand binding.

As the major facilitators of the turnover of organic matter in the marine environment, the ability of heterotrophic bacteria to acquire specific compounds within the diverse range of dissolved organic matter will affect the regeneration of essential nutrients such as iron and carbon. TonB-dependent transporters are a prevalent cellular tool in Gram-negative bacteria that allow a relatively high-molecular-weight fraction of organic matter to be directly accessed. However, these transporters are not well characterized in marine bacteria, limiting our understanding of the flow of specific substrates through the marine microbial loop. Here, we characterize the TonB-dependent transporters responsible for iron and carbon acquisition in a representative marine copiotroph and examine their distribution across the genus Alteromonas . We provide evidence that substrate-specific bioavailability is niche specific, particularly for iron complexes, indicating that transport capacity may serve as a significant control on microbial community dynamics and the resultant cycling of organic matter. Iron is an essential micronutrient for all microbial growth in the marine environment, and in heterotrophic bacteria, iron is tightly linked to carbon metabolism due to its central role as a cofactor in enzymes of the respiratory chain. Here, we present the iron- and carbon-regulated transcriptomes of a representative marine copiotroph, Alteromonas macleodii ATCC 27126, and characterize its cellular transport mechanisms. ATCC 27126 has distinct metabolic responses to iron and carbon limitation and, accordingly, uses distinct sets of TonB-dependent transporters for the acquisition of iron and carbon. These distinct sets of TonB-dependent transporters were of a similar number, indicating that the diversity of carbon and iron substrates available to ATCC 27126 is of a similar scale. For the first time in a marine bacterium, we have also identified six characteristic inner membrane permeases for the transport of siderophores via an ATPase-independent mechanism. An examination of the distribution of specific TonB-dependent transporters in 31 genomes across the genus Alteromonas points to niche specialization in transport capacity, particularly for iron. We conclude that the substrate-specific bioavailability of both iron and carbon in the marine environment will likely be a key control on the processing of organic matter through the microbial loop. IMPORTANCE As the major facilitators of the turnover of organic matter in the marine environment, the ability of heterotrophic bacteria to acquire specific compounds within the diverse range of dissolved organic matter will affect the regeneration of essential nutrients such as iron and carbon. TonB-dependent transporters are a prevalent cellular tool in Gram-negative bacteria that allow a relatively high-molecular-weight fraction of organic matter to be directly accessed. However, these transporters are not well characterized in marine bacteria, limiting our understanding of the flow of specific substrates through the marine microbial loop. Here, we characterize the TonB-dependent transporters responsible for iron and carbon acquisition in a representative marine copiotroph and examine their distribution across the genus Alteromonas . We provide evidence that substrate-specific bioavailability is niche specific, particularly for iron complexes, indicating that transport capacity may serve as a significant control on microbial community dynamics and the resultant cycling of organic matter.

In August 2010, iron (Fe) and Fe and copper (Cu) addition incubation experiments were conducted at two low Fe stations (P20 and P26) along Line P, off the western coast of British Columbia, to investigate Cu physiology in Fe- and Fe-light co-limited phytoplankton. Chlorophyll a concentrations ([Chl a]), maximum variable fluorescence yield (F v/F m), and Fe uptake rates by the Cu-dependent high-affinity Fe transport system (HAFeTS) were measured. Additions of Fe resulted in an increase in [Chl a] and F v/F m at both stations compared with the controls, regardless of light availability, and confirmed that the phytoplankton communities were Fe-limited. Uptake of Fe by the HAFeTS in both incubations increased with the addition of Fe, and likely reflects luxury Fe uptake and storage. While the in situ inorganic Cu concentrations were similar to those that can induce Cu-limitation in laboratory cultures, increasing Cu availability had no effect on biomass accumulation during both incubations, regardless of Fe availability or light regime. At P26, additions of 1 nmol L−1 CuSO₄ resulted in a short-term increase in F v/F m of the phytoplankton community, and an increase in Fe uptake rates by large phytoplankton (>5 μm), but only when light was not limiting. These data confirm a complex interaction between light, Fe and Cu physiology in indigenous phytoplankton communities, and suggest that these interactions may be both spatially heterogeneous and different for different phytoplankton size classes.

The distributions of dissolved iron and organic iron-binding ligands were examined in water column profiles and deckboard incubation experiments in the southern California Current System (sCCS) along a transition from coastal to semi-oligotrophic waters. Analysis of the iron-binding ligand pool by competitive ligand exchange-adsorptive cathodic stripping voltammetry (CLE-ACSV) using multiple analytical windows (MAWs) revealed three classes of iron-binding ligands present throughout the water column (L1-L3), whose distributions closely matched those of dissolved iron and nitrate. Despite significant biogeochemical gradients, ligand profiles were similar between stations, with surface minima in strong ligands (L1 and L2), and relatively constant concentrations of weaker ligands (L3) down to 500 m. A phytoplankton grow-out incubation, initiated from an iron-limited water mass, showed dynamic temporal cycling of iron-binding ligands. A biological iron model was able to capture the patterns of the strong ligands in the grow-out incubation relatively well with only the microbial community as a biological source. An experiment focused on remineralization of particulate organic matter showed production of both strong and weak iron-binding ligands by the heterotrophic community, supporting a mechanism for in-situ production of both strong and weak iron-binding ligands in the subsurface water column. Photochemical experiments showed a variable influence of sunlight on the degradation of natural iron-binding ligands, providing some evidence to explain differences in surface ligand concentrations between stations. Patterns in ligand distributions between profiles and in the incubation experiments were primarily related to macronutrient concentrations, suggesting microbial remineralization processes might dominate on longer time-scales over short-term changes associated with photochemistry or phytoplankton growth.

The roles of iron and light as limiting and colimiting factors for phytoplankton growth in subsurface chlorophyll maxima (SCMs) were investigated in mesotrophic to oligotrophic waters of the Southern California Bight and the eastern tropical North Pacific using microcosm manipulation experiments. Phytoplankton responses indicative of iron-light colimitation were found at several SCMs underlying macronutrient-limited surface waters in the eastern Pacific. Iron additions led to a shift in the size and taxonomic structure of the phytoplankton community, where large diatoms dominated what was formerly a diverse community of relatively small phytoplankton. The strongest and most ubiquitous responses of diatoms to iron addition were found under elevated light conditions, indicating that iron availability may have the greatest potential to affect SCM phytoplankton communities when light levels increase rapidly, such as during eddy events or with strong internal waves. The results show that iron influences phytoplankton community structure at SCMs, which would have consequences for nutrient cycling and carbon export within the lower euphotic zone.