Regulation of Algal Blooms in Antarctic Shelf Waters by the Release of Iron From Melting Sea Ice

During summer 1995-96, we measured iron in the water column and conducted iron-enrichment bottle-incubation experiments at a station in the central Ross Sea (76°30\u27S, 170°40\u27W), first, in the presence of melting sea ice, and 17 days later, in ice-free conditions. We observed a striking temporal change in mixed-layer dissolved iron concentrations at this station, from 0.72-2.3 nM with sea ice present, to 0.16-0.17 nM in ice-free conditions. These changes were accompanied doubling of algal (diatom) biomass. Our incubation experiments suggest that conditions were iron-replete in the presence of sea ice, and iron-deficient in the absence of sea ice. We surmise that bioavailability iron was released into seawater from the melting sea ice, stimulating phytoplankton production and the biological removal of dissolved iron from the mixed layer, until iron-limited conditions developed. These observations suggest that the episodic release of bio-available iron from melting sea ice is an important factor regulating phytoplankton production, particularly ice-edge blooms, in seasonally ice-covered Antarctic waters

Iron and Manganese in the Ross Sea, Antarctica: Seasonal Iron Limitation in Antarctic Shelf Waters

Dissolved iron and manganese and total dissolvable iron were measured in water column samples from the Polynya Region of the southern Ross Sea in cruises in Nov.-Dec. 1994 (spring 1994) and Dec. 1995 to Jan. 1996 (summer 1995). Iron and manganese addition bottle incubation experiments were also performed on these cruises to assess the nutritional sufficiency of ambient iron and manganese concentrations for phytoplankton growth. Generally high dissolved iron concentrations (\u3e0.5 nM) and relatively complex iron and manganese vertical profiles were obtained in spring 1994 vs. summer 1995. Dissolved iron concentrations in the upper water column averaged 1.0 nM in spring 1994 and 0.23 nM in summer 1995, excluding 2 stations where concentrations exceeding 1 nM were attributed to inputs from melting sea ice. The Observed differences in the iron and manganese distribution between spring 1994 and summer 1995 were attributed to seasonal decreases in bottom water upwelling and sea ice melting, which supplied these metals to the upper water column, combined with the cumulative removal of iron and manganese from the water column throughout the spring and summer, due to biological uptake, vertical export, and scavenging by suspended and sinking particles. Results of metal addition bottle incubation experiments indicated that ambient dissolved Iron concentrations were adequate for phytoplankton growth requirements in spring and early summer, when algal production is highest and Phaeocystis antarctica dominates the algal community, whereas low dissolved Iron concentrations limited algal community growth later in the summer, except in stratified, Iron enriched water near melting sea ice, where diatoms are able to bloom. Observations and inferred seasonal distribution of P. antarctica and diatoms in this water suggested that iron availability and vertical mixing (i.e., irradiance) exert the primary controls on phytoplankton growth and community structure in the southern Ross Sea in spring and summer

Influence of Irradiance and Iron on the Growth of Colonial Phaeocystic antarctica: Implications for Seasonal Bloom Dynamics in the Ross Sea, Antarctica

Laboratory culture experiments were used to investigate the growth rate of colonial Phaeocystis anarctica as a function of irradiance and dissolved iron concentration. The experiments were conducted with a P. antarctica strain isolated from the southern Ross Sea, Antarctica, and made use of natural, low-iron (P. antarctica attained an average maximum cell-specific growth rate of 0.37 d-1at an irradiance of 68 μE m-2s-1, above which growth rates decreased to 0.27 d-1 at an irradiance of 314 μE m-2s-1. The dependence of growth rate on ambient dissolved iron concentration was examined in dose-response type bioassay experiments using realistic subnanomolar additions of dissolved iron. The experimental results indicate significant changes in the iron requirements for growth of colonial P. antarctica as a function of irradiance, with our estimates of the half-saturation constant for growth with respect to dissolved iron (Kμ) ranging from 0.26 nM at ~20 μE m-2s-1, to 0.045 nM at similar to 40 μE m-2s-1 and to 0.19 nM at ~ 90 μE m-2 s-1. We interpret these variations in K, as reflecting an increase in the cellular iron requirements of colonial P. antarctica at suboptimal and supraoptimal irradiance, such that the cells require higher ambient dissolved iron concentrations to attain maximum growth rates under Such irradiance conditions. The experiments also provide evidence of a relationship between iron availability and the relative proportion of colonial versus solitary P. antarctica cells, whereby the colonial form appears to be favored by higher dissolved iron concentrations. Our experimental results suggest that the initiation and termination of colonial P. antarctica blooms in the Ross Sea are determined by the combined effects of irradiance-driven changes in cellular iron requirements and a seasonal decrease in dissolved iron availability

Temporal and spatial patterns in the Ross Sea: Phytoplankton biomass, elemental composition, productivity and growth rates

The temporal and spatial patterns of phytoplankton biomass, productivity, and particulate matter composition in the Ross Sea were assessed during cruises in January 1990 and February 1992. Biomass and primary productivity in the southern Ross Sea were greatest during mid-January, with surface chlorophyll concentrations, particulate organic carbon levels, and integrated primary productivity averaging 4.9 Ixg L \u27l, 0.54 mg L-•, and 2.63 g C m • d \u27•, respectively. Comparable mean concentrations and rates for February were 1.1 Ixg L \u27l, 0.29 mg L \u27l, and 0.78 g C m \u27•- d \u27• (decreases of 76, 46, and 70%, respectively), indicative of the scale of temporal changes. A distinct south-north transition also was observed both in productivity and phytoplankton biomass, with the lowest values occurring in the northern Ross Sea. East-west gradients in phytoplankton biomass and composition occurred within the southern Ross Sea. The areal productivity of the Ross Sea ranged from 0.15 to 2.85 g C m • d -• and is among the highest found in the entire Antarctic. Carbon:chlorophyll ratios were uniformly high but were highest (150) in 1990 in the diatom-dominated western Ross Sea. Surface growth rates were modest, averaging less than 0.2 day \u27• during both seasons. We hypothesize that the marked seasonality in the region provides an environment in which net growth rates, although slow, are maximized through low loss rates and which allows biomass to accumulate in the surface layer. Furthermore, the temporal variations are quantitatively similar to the observed spatial variations. Therefore the dominant determinant of phytoplankton biomass and productivity at any one point on the Ross Sea continental shelf is the stage of the seasonal growth cycle

Targeted metaproteomics : detecting sub-species level protein biomarkers in the vast oceanic microbial metaproteome

Author Posting. © The Author(s), 2015. This is the author's version of the work. It is posted here by permission of John Wiley & Sons for personal use, not for redistribution. The definitive version was published in Proteomics 15 (2015): 3521-3531, doi:10.1002/pmic.201400630.Proteomics has great potential for studies of marine microbial biogeochemistry, yet high microbial diversity in many locales presents us with unique challenges. We addressed this challenge with a targeted metaproteomics workflow for NtcA and P-II, two nitrogen regulatory proteins, and demonstrated its application for cyanobacterial taxa within microbial samples from the Central Pacific Ocean. Using METATRYP, an open-source Python toolkit, we examined the number of shared (redundant) tryptic peptides in representative marine microbes, with the number of tryptic peptides shared between different species typically being 1% or less. The related cyanobacteria Prochlorococcus and Synechococcus shared an average of 4.8+1.9% of their tryptic peptides, while shared intraspecies peptides were higher, 13+15% shared peptides between 12 Prochlorococcus genomes. An NtcA peptide was found to target multiple cyanobacteria species, whereas a P-II peptide showed specificity to the high-light Prochlorococcus ecotype. Distributions of NtcA and P-II in the Central Pacific Ocean were similar except at the Equator likely due to differential nitrogen stress responses between Prochlorococcus and Synechococcus. The number of unique tryptic peptides coded for within three combined oceanic microbial metagenomes was estimated to be ~4x107, 1000-fold larger than an individual microbial proteome and 27-fold larger than the human proteome, yet still 20 orders of magnitude lower than the peptide diversity possible in all protein space, implying that peptide mapping algorithms should be able to withstand the added level of complexity in metaproteomic samples.This research was funded by the Gordon and Betty Moore Foundation and the US National Science Foundation under grant numbers 3782, 3934, OCE-1260233, OCE-1233261, OCE-1220484, OCE-1333212 and OCE-1155566, and the Center for Microbial Oceanography Research and Education (C-MORE).2016-06-1

A seasonal study of dissolved cobalt in the Ross Sea, Antarctica : micronutrient behavior, absence of scavenging, and relationships with Zn, Cd, and P

© The Authors, 2010. This article is distributed under the terms of the Creative Commons Attribution 3.0 License. The definitive version was published in Biogeosciences 7 (2010): 4059-4082, doi:10.5194/bg-7-4059-2010.We report the distribution of cobalt (Co) in the Ross Sea polynya during austral summer 2005–2006 and the following austral spring 2006. The vertical distribution of total dissolved Co (dCo) was similar to soluble reactive phosphate (PO43−), with dCo and PO43− showing a significant correlation throughout the water column (r2 = 0.87, 164 samples). A strong seasonal signal for dCo was observed, with most spring samples having concentrations ranging from ~45–85 pM, whereas summer dCo values were depleted below these levels by biological activity. Surface transect data from the summer cruise revealed concentrations at the low range of this seasonal variability (~30 pM dCo), with concentrations as low as 20 pM observed in some regions where PO43− was depleted to ~0.1 μM. Both complexed Co, defined as the fraction of dCo bound by strong organic ligands, and labile Co, defined as the fraction of dCo not bound by these ligands, were typically observed in significant concentrations throughout the water column. This contrasts the depletion of labile Co observed in the euphotic zone of other ocean regions, suggesting a much higher bioavailability for Co in the Ross Sea. An ecological stoichiometry of 37.6 μmol Co:mol−1 PO43− calculated from dissolved concentrations was similar to values observed in the subarctic Pacific, but approximately tenfold lower than values in the Eastern Tropical Pacific and Equatorial Atlantic. The ecological stoichiometries for dissolved Co and Zn suggest a greater overall use of Zn relative to Co in the shallow waters of the Ross Sea, with a Co:PO43−/Zn:PO43− ratio of 1:17. Comparison of these observed stoichiometries with values estimated in culture studies suggests that Zn is a key micronutrient that likely influences phytoplankton diversity in the Ross Sea. In contrast, the observed ecological stoichiometries for Co were below values necessary for the growth of eukaryotic phytoplankton in laboratory culture experiments conducted in the absence of added zinc, implying the need for significant Zn nutrition in the Zn-Co cambialistic enzymes. The lack of an obvious kink in the dissolved Co:PO43− relationship was in contrast to Zn:PO43− and Cd:PO43− kinks previously observed in the Ross Sea. An excess uptake mechanism for kink formation is proposed as a major driver of Cd:PO43− kinks, where Zn and Cd uptake in excess of that needed for optimal growth occurs at the base of the euphotic zone, and no clear Co kink occurs because its abundances are too low for excess uptake. An unusual characteristic of Co geochemistry in the Ross Sea is an apparent lack of Co scavenging processes, as inferred from the absence of dCo removal below the euphotic zone. We hypothesize that this vertical distribution reflects a low rate of Co scavenging by Mn oxidizing bacteria, perhaps due to Mn scarcity, relative to the timescale of the annual deep winter mixing in the Ross Sea. Thus Co exhibits nutrient-like behavior in the Ross Sea, in contrast to its hybrid-type behavior in other ocean regions, with implications for the possibility of increased marine Co inventories and utility as a paleooceanographic proxy.This research was supported by the US National Science Foundation through research grants (OPP-0440840, OPP-0338097, OPP-0732665, OCE-0452883, OCE-0752991, OCE-0928414)

Thaumarchaeal ecotype distributions across the equatorial Pacific Ocean and their potential roles in nitrification and sinking flux attenuation

© The Author(s), 2017. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Limnology and Oceanography 62 (2017): 1984–2003, doi:10.1002/lno.10547.Thaumarchaea are among the most abundant microbial groups in the ocean, but controls on their abundance and the distribution and metabolic potential of different subpopulations are poorly constrained. Here, two ecotypes of ammonia-oxidizing thaumarchaea were quantified using ammonia monooxygenase (amoA) genes across the equatorial Pacific Ocean. The shallow, or water column “A” (WCA), ecotype was the most abundant ecotype at the depths of maximum nitrification rates, and its abundance correlated with other biogeochemical indicators of remineralization such as NO3 : Si and total Hg. Metagenomes contained thaumarchaeal genes encoding for the catalytic subunit of the urease enzyme (ureC) at all depths, suggesting that members of both WCA and the deep, water column “B” (WCB) ecotypes may contain ureC. Coupled urea hydrolysis-ammonia oxidation rates were similar to ammonia oxidation rates alone, suggesting that urea could be an important source of ammonia for mesopelagic ammonia oxidizers. Potential inducement of metal limitation of both ammonia oxidation and urea hydrolysis was demonstrated via additions of a strong metal chelator. The water column inventory of WCA was correlated with the depth-integrated abundance of WCB, with both likely controlled by the flux of sinking particulate organic matter, providing strong evidence of vertical connectivity between the ecotypes. Further, depth-integrated amoA gene abundance and nitrification rates were correlated with particulate organic nitrogen flux measured by contemporaneously deployed sediment traps. Together, the results refine our understanding of the controls on thaumarchaeal distributions in the ocean, and provide new insights on the relationship between material flux and microbial communities in the mesopelagic.United States National Science Foundation (NSF) Grant Numbers: OCE-1260006, OCE-1031271, OCE-1337780, OCE-1259994; University of Maryland Center for Environmental Science (UMCES); JGI Community Sequencing Project 133

A Seasonal Study of Dissolved Cobalt in the Ross Sea, Antarctica: Micronutrient Behavior, Absence of Scavenging, and Relationships with Zd, Cd, and P.

We report the distribution of cobalt (Co) in the Ross Sea polynya during austral summer 2005-2006 and the following austral spring 2006. The vertical distribution of total dissolved Co (dCo) was similar to soluble reactive phosphate (PO(4)(3-)), with dCo and PO(4)(3-) showing a significant correlation throughout the water column (r(2) = 0.87, 164 samples). A strong seasonal signal for dCo was observed, with most spring samples having concentrations ranging from similar to ~ 45-85 pM, whereas summer dCo values were depleted below these levels by biological activity. Surface transect data from the summer cruise revealed concentrations at the low range of this seasonal variability (similar to ~ 30pM dCo), with concentrations as low as 20pM observed in some regions where PO(4)(3-) was depleted to similar to 0.1 mu M. Both complexed Co, defined as the fraction of dCo bound by strong organic ligands, and labile Co, defined as the fraction of dCo not bound by these ligands, were typically observed in significant concentrations throughout the water column. This contrasts the depletion of labile Co observed in the euphotic zone of other ocean regions, suggesting a much higher bioavailability for Co in the Ross Sea. An ecological stoichiometry of 37.6 mu mol Co: mol(-1) PO(4)(3-) calculated from dissolved concentrations was similar to values observed in the subarctic Pacific, but approximately tenfold lower than values in the Eastern Tropical Pacific and Equatorial Atlantic. The ecological stoichiometries for dissolved Co and Zn suggest a greater overall use of Zn relative to Co in the shallow waters of the Ross Sea, with a Co: PO(4)(3-) / Zn: PO(4)(3-) ratio of 1:17. Comparison of these observed stoichiometries with values estimated in culture studies suggests that Zn is a key micronutrient that likely influences phytoplankton diversity in the Ross Sea. In contrast, the observed ecological stoichiometries for Co were below values necessary for the growth of eukaryotic phytoplankton in laboratory culture experiments conducted in the absence of added zinc, implying the need for significant Zn nutrition in the Zn-Co cambialistic enzymes. The lack of an obvious kink in the dissolved Co: PO(4)(3-) relationship was in contrast to Zn: PO(4)(3-) and Cd: wPO(4)(3-) kinks previously observed in the Ross Sea. An excess uptake mechanism for kink formation is proposed as a major driver of Cd: PO(4)(3-) kinks, where Zn and Cd uptake in excess of that needed for optimal growth occurs at the base of the euphotic zone, and no clear Co kink occurs because its abundances are too low for excess uptake. An unusual characteristic of Co geochemistry in the Ross Sea is an apparent lack of Co scavenging processes, as inferred from the absence of dCo removal below the euphotic zone. We hypothesize that this vertical distribution reflects a low rate of Co scavenging by Mn oxidizing bacteria, perhaps due to Mn scarcity, relative to the timescale of the annual deep winter mixing in the Ross Sea. Thus Co exhibits nutrient-like behavior in the Ross Sea, in contrast to its hybrid-type behavior in other ocean regions, with implications for the possibility of increased marine Co inventories and utility as a paleooceanographic proxy

Biogeochemical and ecological variability during the late summer–early autumn transition at an ice‐floe drift station in the Central Arctic Ocean

As the annual expanse of Arctic summer ice‐cover steadily decreases, concomitant biogeochemical and ecological changes in this region are likely to occur. Because the Central Arctic Ocean is often nutrient and light limited, it is essential to understand how environmental changes will affect productivity, phytoplankton species composition, and ensuing changes in biogeochemistry in the region. During the transition from late summer to early autumn, water column sampling of various biogeochemical parameters was conducted along an ice‐floe drift station near the North Pole. Our results show that as the upper water column stratification weakened during the late summer–early autumn transition, nutrient concentrations, particulate dimethylsulfoniopropionate (DMSPp) levels, photosynthetic efficiency, and biological productivity, as estimated by ΔO2/Ar ratios, all decreased. Chemotaxonomic (CHEMTAX) analysis of phytoplankton pigments revealed a taxonomically diverse picoautotrophic community, with chlorophyll (Chl) c3‐containing flagellates and the prasinophyte, Pyramimonas spp., as the most abundant groups, comprising ~ 30% and 20% of the total Chl a (TChl a) biomass, respectively. In contrast to previous studies, the picoprasinophyte, Micromonas spp., represented only 5% to 10% of the TChl a biomass. Of the nine taxonomic groups identified, DMSPp was most closely associated with Pyramimonas spp., a Chl b‐containing species not usually considered a high DMSP producer. As the extent and duration of open, ice‐free waters in the Central Arctic Ocean progressively increases, we suggest that enhanced light transmission could potentially expand the ecological niche of Pyramimonas spp. in the region

Iron limitation of a springtime bacterial and phytoplankton community in the Ross Sea : implications for vitamin B12 nutrition

© The Author(s), 2011. This is an open-access article subject to a non-exclusive license between the authors and Frontiers Media SA, which permits use, distribution and reproduction in other forums. The definitive version was published in Frontiers in Microbiology 2 (2011): 160, doi:10.3389/fmicb.2011.00160.The Ross Sea is home to some of the largest phytoplankton blooms in the Southern Ocean. Primary production in this system has previously been shown to be iron limited in the summer and periodically iron and vitamin B12 colimited. In this study, we examined trace metal limitation of biological activity in the Ross Sea in the austral spring and considered possible implications for vitamin B12 nutrition. Bottle incubation experiments demonstrated that iron limited phytoplankton growth in the austral spring while B12, cobalt, and zinc did not. This is the first demonstration of iron limitation in a Phaeocystis antarctica-dominated, early season Ross Sea phytoplankton community. The lack of B12 limitation in this location is consistent with previous Ross Sea studies in the austral summer, wherein vitamin additions did not stimulate P. antarctica growth and B12 was limiting only when bacterial abundance was low. Bottle incubation experiments and a bacterial regrowth experiment also revealed that iron addition directly enhanced bacterial growth. B12 uptake measurements in natural water samples and in an iron fertilized bottle incubation demonstrated that bacteria serve not only as a source for vitamin B12, but also as a significant sink, and that iron additions enhanced B12 uptake rates in phytoplankton but not bacteria. Additionally, vitamin uptake rates did not become saturated upon the addition of up to 95 pM B12. A rapid B12 uptake rate was observed after 13 min, which then decreased to a slower constant uptake rate over the next 52 h. Results from this study highlight the importance of iron availability in limiting early season Ross Sea phytoplankton growth and suggest that rates of vitamin B12 production and consumption may be impacted by iron availability.This research was supported by NSF grants OCE-0752291, OPP-0440840, OPP-0338097, OPP-0338164, ANT-0732665, OCE-0452883, and OCE-1031271, the Center for Microbial Oceanography Research and Education (CMORE) and a National Science Foundation (NSF) Graduate Research Fellowship (2007037200) and an Environmental Protection Agency STAR Fellowship to EMB (F6E20324)