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Phytoplankton iron contents (i.e., quotas) directly link biogeochemical cycles of iron and carbon and drive patterns of nutrient limitation, recycling, and export. Ocean biogeochemical models typically assume that iron quotas are either static or controlled by dissolved iron availability. We measured iron quotas in phytoplankton communities across nutrient gradients in the Pacific Ocean and found that quotas diverged significantly in taxon‐specific ways from laboratory‐derived predictions. Iron quotas varied 40‐fold across nutrient gradients, and nitrogen‐limitation allowed diatoms to accumulate fivefold more iron than co‐occurring flagellates even under low iron availability. Modeling indicates such “luxury” uptake is common in large regions of the low‐iron Pacific Ocean. Among diatoms, both pennate and centric genera accumulated luxury iron, but the cosmopolitan pennate genus Pseudo‐nitzschia maintained iron quotas 10‐fold higher than co‐occurring centric diatoms, likely due to enhanced iron storage. Biogeochemical models should account for taxonomic and macronutrient controls on phytoplankton iron quotas.

Significance The vertical distribution of phytoplankton cells and chlorophyll concentrations throughout the sunlit water column is rarely uniform. In many ocean regions, chlorophyll concentrations peak in distinct and persistent layers deep below the surface called subsurface chlorophyll maximum layers (SCMLs). SCML formation is hypothesized to reflect the consequences of phytoplankton light/macronutrient colimitation, behavior, and/or photoacclimation. We discovered unexpectedly persistent and widespread phytoplankton iron limitation and iron/light colimitation in SCMLs of the California Current and at the edge of the North Pacific Subtropical Gyre using shipboard incubations, metatranscriptomics, and biogeochemical proxies. These results suggest that interactions and feedbacks between iron and light availability play an important and previously unrecognized role in controlling the productivity and biogeochemical dynamics of SCMLs., Subsurface chlorophyll maximum layers (SCMLs) are nearly ubiquitous in stratified water columns and exist at horizontal scales ranging from the submesoscale to the extent of oligotrophic gyres. These layers of heightened chlorophyll and/or phytoplankton concentrations are generally thought to be a consequence of a balance between light energy from above and a limiting nutrient flux from below, typically nitrate (NO3). Here we present multiple lines of evidence demonstrating that iron (Fe) limits or with light colimits phytoplankton communities in SCMLs along a primary productivity gradient from coastal to oligotrophic offshore waters in the southern California Current ecosystem. SCML phytoplankton responded markedly to added Fe or Fe/light in experimental incubations and transcripts of diatom and picoeukaryote Fe stress genes were strikingly abundant in SCML metatranscriptomes. Using a biogeochemical proxy with data from a 40-y time series, we find that diatoms growing in California Current SCMLs are persistently Fe deficient during the spring and summer growing season. We also find that the spatial extent of Fe deficiency within California Current SCMLs has significantly increased over the last 25 y in line with a regional climate index. Finally, we show that diatom Fe deficiency may be common in the subsurface of major upwelling zones worldwide. Our results have important implications for our understanding of the biogeochemical consequences of marine SCML formation and maintenance.

Tidal Spartina-dominated saltmarshes and estuaries on the Southeast US coast are global hotspots of productivity. In coastal Georgia, tidal amplitudes and saltmarsh productivity are the highest along the Southeast US coast. Coastal Georgia is characterized by a humid subtropical seasonal climate, and inter-annual variability in precipitation, and freshwater discharge. The 2012–2013 timeframe encompassed contrasting levels of discharge for the Savannah River, a major Georgia river, with a 4.3-fold greater discharge in summer 2013 relative to summer 2012. In situ measurements of temperature, salinity, precipitation and Secchi depth, and water samples were collected weekly at high tide throughout 2012 and 2013 from the Skidaway River Estuary, a tidal saltmarsh-dominated estuary in coastal Georgia influenced by Savannah River hydrology. The effects of elevated discharge on the seasonal trends of water column components were evaluated. The shift from low discharge (2012) to high discharge (2013) led to decreased salinity in summer 2013, but no significant increases in inorganic nutrient (NH4, NOx, SiO2 and PO4) concentrations. Dissolved inorganic carbon (DIC) concentrations decreased, and DIC stable isotopic signatures (δ13C-DIC values) were depleted in summer 2013 relative to summer 2012. In 2013 dissolved organic carbon (DOC) concentrations, chromophoric and fluorescent dissolved organic matter (DOM: CDOM, FDOM) intensities, specific UV-absorbance (SUVA254) and relative humic-like fluorescence were all higher than in 2012, indicating that, as discharge increased in 2013, estuarine water became enriched in terrigenous DOM. Secchi depth and particulate organic carbon (POC) and nitrogen (PON) concentrations displayed clear seasonal patterns that were not significantly altered by discharge. However, δ13C-POC and δ15N-PON isotopic signatures indicated higher terrigenous contributions at elevated discharge. Discharge influenced cyanobacterial composition, but did not affect total abundance of phytoplankton (

Diatoms are prominent eukaryotic phytoplankton despite being limited by the micronutrient iron in vast expanses of the ocean. As iron inputs are often sporadic, diatoms have evolved mechanisms such as the ability to store iron that enable them to bloom when iron is resupplied and then persist when low iron levels are reinstated. Two iron storage mechanisms have been previously described: the protein ferritin and vacuolar storage. To investigate the ecological role of these mechanisms among diatoms, iron addition and removal incubations were conducted using natural phytoplankton communities from varying iron environments. We show that among the predominant diatoms, Pseudo-nitzschia were favored by iron removal and displayed unique ferritin expression consistent with a long-term storage function. Meanwhile, Chaetoceros and Thalassiosira gene expression aligned with vacuolar storage mechanisms. Pseudo-nitzschia also showed exceptionally high iron storage under steady-state high and low iron conditions, as well as following iron resupply to iron-limited cells. We propose that bloom-forming diatoms use different iron storage mechanisms and that ferritin utilization may provide an advantage in areas of prolonged iron limitation with pulsed iron inputs. As iron distributions and availability change, this speculated ferritin-linked advantage may result in shifts in diatom community composition that can alter marine ecosystems and biogeochemical cycles.

Transitions in community genomic features and biogeochemical processes were examined in surface and subsurface chlorophyll maximum (SCM) microbial communities across a trophic gradient from mesotrophic waters near San Diego, California to the oligotrophic Pacific. Transect end points contrasted in thermocline depth, rates of nitrogen and CO2 uptake, new production and SCM light intensity. Relative to surface waters, bacterial SCM communities displayed greater genetic diversity and enrichment in putative sulfur oxidizers, multiple actinomycetes, low-light-adapted Prochlorococcus and cell-associated viruses. Metagenomic coverage was not correlated with transcriptional activity for several key taxa within Bacteria. Low-light-adapted Prochlorococcus, Synechococcus, and low abundance gamma-proteobacteria enriched in the>3.0-μm size fraction contributed disproportionally to global transcription. The abundance of these groups also correlated with community functions, such as primary production or nitrate uptake. In contrast, many of the most abundant bacterioplankton, including SAR11, SAR86, SAR112 and high-light-adapted Prochlorococcus, exhibited low levels of transcriptional activity and were uncorrelated with rate processes. Eukaryotes such as Haptophytes and non-photosynthetic Aveolates were prevalent in surface samples while Mamielles and Pelagophytes dominated the SCM. Metatranscriptomes generated with ribosomal RNA-depleted mRNA (total mRNA) coupled to in vitro polyadenylation compared with polyA-enriched mRNA revealed a trade-off in detection eukaryotic organelle and eukaryotic nuclear origin transcripts, respectively. Gene expression profiles of SCM eukaryote populations, highly similar in sequence identity to the model pelagophyte Pelagomonas sp. CCMP1756, suggest that pelagophytes are responsible for a majority of nitrate assimilation within the SCM.

The goal of the current study was to test a scoring matrix for evaluating and quantifying the difficulty of use of consumer packaging for an older adult population. Scores range from 0 to 100; the lower the score, the less usable the packaging design. Twelve older adults (9 females, 3 males; Mage = 69.5) completed an online survey in which they evaluated 15 everyday packaging types on frequency of use and difficulty of the functional tasks required for use (e.g., pick up and carry, remove seal, open, dispense, close). Blister packaging scored the lowest ( M = 71.5) and could benefit the most from redesign. Microwave meal packaging scored the highest ( M = 85.8), but this was the only score computed without all participants; only five participants reported using microwave meals. Pull tab cans scored second highest with a mean score of 84.2. In this pilot study, we are laying the groundwork for creating a tool to systematically evaluate and quantify the usability of all packaging types.

The increasing proportion of the population that is over age 65 stimulates the need for more technologies that can be safely and easily used by older adults to allow them to continue living in their homes. Yet, even designers who know typical age-related declines may not understand specifically how the declines affect interaction with everyday technologies to complete activities of daily living. Thus, technologies may not be effectively designed to accommodate these declines. Through demonstration, participants will develop this experience by completing activities typical of older adults while wearing tools that simulate typical sensory and motor declines. Facilitators will encourage discussion of the specific difficulties experienced by participants with other observers, highlighting features that may facilitate or hinder use given the simulated disability. Highlighting limitations and task difficulties that may be experienced through use of simulation tools will encourage a better understanding of universal design criteria that will be more inclusive of the overall population.

Trichodesmium colonies contain an abundant microbial consortium that is likely to play a role in nutrient cycling within the colony. This study used laboratory cultures of Trichodesmium and two genome‐sequenced strains of bacteria typical of Trichodesmium‐associated microbes to develop an understanding of the cycling of iron, a potentially limiting micronutrient, within Trichodesmium colonies. We found that the ferric siderophores desferrioxamine B and aerobactin were not readily bioavailable to Trichodesmium, relative to ferric chloride or citrate‐associated iron. In contrast, the representative bacterial strains we studied were able to acquire iron from all of the iron sources, implying that naturally occurring Trichodesmium‐associated bacteria may be capable of utilizing a more diverse array of iron sources than Trichodesmium. From the organism‐specific uptake data collected in this study, a theoretical Trichodesmium colony was designed to model whole colony iron uptake. The bacteria accounted for most (> 70%) of the iron acquired by the colony, highlighting the importance of determining organism‐specific uptake in a complex environment. Our findings suggest that, although they may share the same micro‐environment, Trichodesmium and its colony‐associated microbial cohort may differ substantially in terms of iron acquisition strategy.

Pathogenic gram-negative bacteria use specific receptors to transport ferric siderophore complexes across their outer membrane during iron (Fe)-limited growth. Receptors such as these have not yet been characterized in oceanic heterotrophic bacteria. We examined four species of g-proteobacteria for the presence of Fe-siderophore receptors with the use of a nondenaturing polyacrylamide gel electrophoresis binding assay and the siderophore ferrioxamine B (FB) labeled with 55 Fe. Small-subunit rRNA sequence analysis assigned these bacteria to the genera Pseudoalteromonas and Alteromonas. Two oceanic species, Pseudoalteromonas haloplanktis (Neptune) and Alteromonas macleodii (Jul88), which were shown previously to transport and assimilate Fe bound to FB during growth, synthesized an outer-membrane FB receptor under Fe-limiting conditions. Only low concentrations of the receptors were detected in these bacteria when they were grown with high concentrations of Fe. The FB receptor of P. haloplanktis (Neptune) had an apparent molecular mass of 79 kDa and an externally oriented binding site. The molecular mass of the receptor of A. macleodii (Jul88) was 100 kDa. No FB receptors were detected by our methods in two coastal species, Pseudoalteromonas rubra(LMG1) and Pseudoalteromonas piscicida(PWF3). P. haloplanktis (Neptune) and A. macleodii (Jul88) also bound 55 Fe-ferrichrome, a trihydroxamate siderophore like FB. Binding assays conducted with 115 nmol L 2 15 5 Fe-FB in the presence of increasing concentrations of desferrioxamine B showed a progressive decrease in the amount of 55 Fe-FB bound by the receptor protein, suggesting strong affinity of the receptor for the Fe-free siderophore. Our results provide the first demonstration of Fe-siderophore receptors in oceanic heterotrophic bacteria. Heterotrophic bacteria have evolved Fe(III) transport systems that enable them to grow in environments containing extraordinarily low concentrations of Fe. When Fe is scarce, many of these organisms excrete low‐molecular mass Febinding compounds, called siderophores, that bind free Fe(III) in solution. The Fe-siderophore complex attaches to its cognate receptor on the outer membrane of the cell and is subsequently internalized (Braun and Killman 1999; Ratledge and Dover 2000). Some bacterial receptors recognize more than one siderophore, not necessarily of the same structural type (e.g., FhuE of E. coli) (van der Helm 1998). Many bacteria express receptors for siderophores released by other species or for Fe sources contained in their hosts (Braun et 1

The closely related cyanobacteria Synechococcus and Prochlorococcus have different distributions in stratified water columns in the northern Sargasso Sea. The abundance of Synechococcus is relatively uniform with depth, but Prochlorococcuscell numbers are low within shallow mixed layers and high in and below the thermocline. Because free cupric ion (free Cu 21 ) concentrations are high (up to 6 pM) in shallow mixed layers and lower in deeper water, there is an inverse relationship between Prochlorococcus densities and the free Cu 21 concentration. We explored the possibility of a causal underpinning for this relationship by examining the relative copper sensitivities of Prochlorococcus and Synechococcus in cultures and field populations. Prochlorococcus isolates from both the high- and low‐light adapted ecotypes were inhibited at free Cu 21 concentrations that had no effect on Synechococcus. However, the high‐light adapted strains were more copper resistant than their low‐light adapted counterparts. When copper was added to Prochlorococcus from environments where the in situ free Cu 21 was low (in deeply mixed water columns and below the mixed layer in stratified conditions), net growth rates were substantially reduced and cells arrested in the G1 and early S phases of the cell cycle. Prochlorococcus in shallow mixed layers were less sensitive to copper and were probably members of the copper-resistant high‐light adapted ecotype. Synechococcus were relatively copper resistant across a range of environments in the Sargasso Sea. These observations are consistent with our hypothesis that copper plays a role in cyanobacteria ecology in the Sargasso Sea.

Prochlorococcus, a small unicellular cyanobacterium, is an important member of the phytoplankton community in the eastern equatorial Pacific. When these waters were enriched with iron during IronEx II, the chlorophyll per cell and cell size of Prochlorococcus increased, implying that they were iron limited. The extent of this limitation was unclear, however, and the number of Prochlorococcus remained constant. To examine whether cell division rates were stimulated significantly by iron, we used a cell cycle analysis approach to measure them in and out of the Fe-enriched patch and in Fe-enriched bottles. The cell division rate increased from 0.6 to 1.1 d 21 over 6 d of exposure to the elevated iron concentrations in the patch. Cells incubated in bottles with additional iron had rates of 1.4 d 21 or two doublings per day. Prochlorococcus mortality rates, measured independently, nearly doubled after the addition of iron. This matched the increase in the cell division rate and maintained a relatively constant population size. Thus the cell division rates of even the smallest phytoplankton in the equatorial Pacific are significantly iron limited, but biomass is constrained by both iron limitation and microzooplankton grazing. The differential response of individual phytoplankton groups to the addition of iron during IronEx II was at least partially a result of differential mortality rates over the time course of the experiment. How the community would respond to sustained fertilization, however, is not obvious.

Recent unenclosed iron-fertilization experiments in the equatorial Pacific Ocean have shown that phytoplankton biomass can be increased substantially by the addition of iron. Analyses of size-fractionated chlorophyll indicate that much of the increase during the most recent fertilization experiment, IronEx II, occurred in the .10-mm size fraction. We used flow cytometry, combined with taxon-specific pigment measurements by high-performance liquid chromatography (HPLC), to analyze the responses of five different groups of phytoplankton:Prochlorococcus, Synechococcus, ultraplankton, nanoplankton, and pennate diatoms. These results are unique in the suite of measurements from the IronEx studies in that they simultaneously examine individual cell properties, which are grazer independent, and population dynamics, which reflect the net result of growth and grazing. Our results show that the overall increase of chlorophyll a (Chl a) in the patch was due in part to increases in chlorophyll content per cell and in part to increases in cell numbers of specific groups. Cellular fluorescence was stimulated by iron addition in all five groups to a qualitatively similar degree and was correlated with taxon-specific changes in cellular pigments. In terms of net cell growth, however, these groups responded very differently. The groups that dominated the community before the addition of iron increased at most twofold in cell number; Prochlorococcus actually decreased. In contrast, the initially rare pennate diatoms increased 15-fold in number by the peak of the iron-induced bloom. Within 1 week, this differential response led to a dramatic change in the phytoplankton community structure, from one dominated by picoplankton to one dominated by large diatoms. It is not known whether this shift would be sustained over extended periods of fertilization, a response that would ultimately change the structure of the food web.

Iron concentrations in the ocean are low enough to limit the growth of marine microorganisms, which raises questions about the molecular mechanisms these organisms use to acquire iron. Marine bacteria have been shown to produce siderophores to facilitate iron(III) uptake. We describe the structures of a suite of amphiphilic siderophores, named the amphibactins, which are produced by a nearshore isolate, γ Proteobacterium , Vibrio sp. R-10. Each amphibactin has the same Tris-hydroxamate-containing peptidic headgroup composed of three ornithine residues and one serine residue but differs in the acyl appendage, which ranges from C-14 to C-18 and varies in the degree of saturation and hydroxylation. Although amphiphilic siderophores are relatively rare, cell-associated amphiphilic siderophores are even less common. We find that the amphibactins are cell-associated siderophores. As a result of the variation in the nature of the fatty acid appendage and the cellular location of the amphibactins, the membrane partitioning of these siderophores was investigated. The physiological mixture of amphibactins had a range of membrane affinities (3.8 × 10 3 to 8.3 × 10 2 M −1 ) that are larger overall than other amphiphilic siderophores, likely accounting for their cell association. This cell association is likely an important defense against siderophore diffusion in the oceanic environment. The phylogenetic affiliation of Vibrio sp. R-10 is discussed, as well as the observed predominance of amphiphilic siderophores produced by marine bacteria in contrast to those produced by terrestrial bacteria.