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The direct respiration of sinking organic matter by attached bacteria is often invoked as the dominant sink for settling particles in the mesopelagic ocean. However, other processes, such as enzymatic solubilization and mechanical disaggregation, also contribute to particle flux attenuation by transferring organic matter to the water column. Here we use observations from the North Atlantic Ocean, coupled to sensitivity analyses of a simple model, to assess the relative importance of particle-attached microbial respiration compared to the other processes that can degrade sinking particles. The observed carbon fluxes, bacterial production rates, and respiration by water column and particle-attached microbial communities each spanned more than an order of magnitude. Rates of substrate-specific respiration on sinking particle material ranged from 0.007 ± 0.003 to 0.173 ± 0.105 day−1. A comparison of these substrate-specific respiration rates with model results suggested sinking particle material was transferred to the water column by various biological and mechanical processes nearly 3.5 times as fast as it was directly respired. This finding, coupled with strong metabolic demand imposed by measurements of water column respiration (729.3 ± 266.0 mg C m−2 d−1, on average, over the 50 to 150 m depth interval), suggested a large fraction of the organic matter evolved from sinking particles ultimately met its fate through subsequent remineralization in the water column. At three sites, we also measured very low bacterial growth efficiencies and large discrepancies between depth-integrated mesopelagic respiration and carbon inputs.

Shipboard bottle incubation experiments were performed to investigate the effects of iron, light, and silicate on algal production of particulate dimethylsulfoniopropionate (DMSPp) in the Subantarctic Zone (SAZ) south of Tasmania during March 1998. Iron enrichment resulted in threefold to ninefold increases in DMSPp concentrations relative to control treatments, following 7 and 8-day incubation experiments. Additions of Fe and Si preferentially stimulated the growth of lightly-silicified pennate diatoms and siliceous haptophytes, respectively, to which we attribute the increased DMSPp production in the incubation bottles. Both of these algal groups were previously believed to be low DMSPp producers; however, our experimental data suggest that addition of iron and silicate to the low-silicate low-iron waters of the SAZ will result in increased production of DMSPp by lightly silicified diatoms and siliceous haptophytes, respectively. Increased irradiance enhanced DMSPp production in iron-amended treatments with both low (0.5 nM) and high (5 nM) concentrations of added iron. However, the role of light in stimulating DMSPp production was apparently of secondary importance compared to the effects of iron addition. The combination of high irradiance and high iron enrichment produced the highest DMSPp production in the experiments, suggesting that iron and light may have a synergistic effect in limiting algal DMSPp production in subantarctic waters.

The influence of irradiance and iron (Fe) supply on phytoplankton processes was investigated, north (47S, 142E) and south (54S, 142E) of the Subantarctic Front in austral autumn (March 1998). At both sites, resident cells exhibited nutrient stress (Fv/ Fm 0.3). Shipboard perturbation experiments examined two light (mean in situ and elevated) and two Fe (nominally 0.5 and 3 nM) treatments under silicic acid-replete conditions. Mean in situ light levels (derived from incident irradiances, mixed layer depths (MLDs), wind stress, and a published vertical mixing model) differed at the two sites, 25% of incident irradiance I 0 at 47S and 9% I 0 at 54S because of MLDs of 40 (47S) and 90 m (54S), when these stations were occupied. The greater MLD at 54S is reflected by tenfold higher cellular chlorophyll a levels in the resident phytoplankton. In the 47S experiment, chlorophyll a levels increased to 1 gL 1 only in the high-Fe treatments, regardless of irradiance levels, suggesting Fe limitation. This trend was also noted for cell abundances, silica production, and carbon fixation rates. In contrast, in the 54S experiment there were increases in chlorophyll a (to 2 gL 1 ), cell abundances, silica production, and carbon fixation only in the high-light treatments to which Fe had been added, suggesting that Fe and irradiance limit algal growth rates. Irradiance by altering algal Fe quotas is a key determinant of algal growth rate at 54S (when silicic acid levels are nonlimiting); however, because of the integral nature of Fe/light colimitation and the restricted nature of the current data set, it was not possible to ascertain the relative contributions of Fe and irradiance to the control of phytoplankton growth. On the basis of a climatology of summer mean MLD for subantarctic (SA) waters south of Australia the 47 and 54S sites appear to represent minimum and maximum MLDs, where Fe and Fe/ irradiance, respectively, may limit/colimit algal growth. The implications for changes in the factors limiting algal growth with season in SA waters are discussed.

Subantarctic Southern Ocean surface waters in the austral summer and autumn are characterized by high concentrations of nitrate and phosphate but low concentrations of dissolved iron (Fe, ∼0.05 nM) and silicic acid (Si

We investigated the hydrography, nutrients, and dissolved and particulate carbon pools in the western Pacific sector of the Antarctic Circumpolar Current (ACC) during austral summer 1996 to assess the region's role in the carbon cycle. Low fCO2 values along two transects indicated that much of the study area was a sink for atmospheric CO2. The fCO2 values were lowest near the Polar Front (PF) and the Subtropical Front (STF), concomitant with maxima of chlorophyll a and particulate and dissolved organic carbon. The largest biomass accumulations did not occur at fronts, which had high surface geostrophic velocities (20–51 cm s−1), but in relatively low velocity regions near fronts or in an eddy. Thus vertical motion and horizontal advection associated with fronts may have replenished nutrients in surface waters but also dispersed phytoplankton. Although surface waters north of the PF have been characterized as a “high nutrient-low chlorophyll” region, low silicic acid (Si) concentrations (2–4 μM) may limit production of large diatoms and therefore the potential carbon flux. Low concentrations (4–10 μM Si) at depths of winter mixing constrain the level of Si replenishment to surface waters. It has been suggested that an increase in aeolian iron north of the PF may increase primary productivity and carbon export. Our results, however, indicate that while diatom growth and carbon export may be enhanced, the extent ultimately would be limited by the vertical supply of Si. South of the PF, the primary mechanism by which carbon is exported to deep water appears to be through diatom flux. We suggest that north of the PF, particulate and dissolved carbon may be exported primarily to intermediate depths through subduction and diapycnal mixing associated with Subantarctic Mode Water and Antarctic Intermediate Water formation. These physical-biological interactions and Si dynamics should be included in future biogeochemical models to provide a more accurate prediction of carbon flux.

Dissolved iron and manganese and total dissolvable iron were measured in water column samples collected from the polynya region of the southern Ross Sea during cruises in November–December 1994 (spring 1994) and December 1995 to January 1996 (summer 1995). Iron and manganese addition bottle incubation experiments were also performed during these cruises in order to assess the nutritional sufficiency of ambient iron and manganese concentrations for growth of the phytoplankton community. Generally high dissolved iron concentrations (>0.5 nM) and relatively complex iron and manganese vertical profiles were obtained during the spring 1994 cruise, compared with the summer 1995 data. Dissolved iron concentrations in the upper water column averaged 1.0 nM during spring 1994 and 0.23 nM in summer 1995, excluding two stations where concentrations exceeding 1 nM are attributed to inputs from melting sea ice. The observed differences in the distribution of iron and manganese between spring 1994 and summer 1995 are attributed to seasonal decreases in the up welling of bottom waters and melting of sea ice, which supply these metals into 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 the metal addition bottle incubation experiments indicate that ambient dissolved iron concentrations are adequate for phytoplankton growth requirements during the spring and early summer, when algal production is highest and Phaeocystis antarctica dominates the algal community, whereas low dissolved iron concentrations limit algal community growth later in the summer, except in the stratified, iron-enriched waters near melting sea ice, where diatoms are able to bloom. Our observations and the inferred seasonal distributions of P. antarctica and diatoms in these waters suggest that iron availability and vertical mixing (i.e., irradiance) exert the primary controls on phytoplankton growth and community structure in the southern Ross Sea during the spring and summer.

Patterns of nutrient utilization and primary productivity (PP) in late austral spring and early summer in the southwestern Ross Sea were characterized with respect to phytoplankton taxonomic composition, polynya dynamics, and upper ocean hydrography during the 1996–1997 oceanographic program Research on Ocean-Atmosphere Variability and Ecosystem Response in the Ross Sea. Phytoplankton biomass in the upper 150 m of the water column ranged from 40 to 540 mg chlorophyll a (Chl a) m−2, exceeding 200 mg Chl a m−2 everywhere except the extreme northern and eastern boundaries of the Ross Sea polynya. Diatom biomass was greatest in the shallow mixed layers of Terra Nova Bay, while the more deeply mixed waters of the Ross Sea polynya were dominated by Phaeocystis antarctica. Daily production computed from the disappearance of NO3 (1.14 g C m−2 d−1) and total dissolved inorganic carbon (TDIC, 1.29 g C m−2 d−1) is consistent with estimates made from an algorithm forced with satellite measurements of Chl a (1.25 g C m−2 d−1) and from measurements of 14C uptake (1.33 g C m−2 d−1). Phytoplankton PP in the Ross Sea averaged 100 g C m−2 yr−1 during 1996–1997. Despite the early formation of the Terra Nova Bay polynya the diatom bloom there did not reach its peak PP until middle to late January 1997 (most likely because of more intense wind mixing in November), ∼6 weeks after the P. antarctica bloom in the Ross Sea polynya had reached the same stage of development. From 70 to 100% of the C and N deficits in the upper 150 m could be accounted for by particulate organic matter, indicating that there had been little dissolved organic matter production or export of particulate material prior to our cruise. This suggests that early in the season, PP and zooplankton grazing are decoupled in the southwestern Ross Sea. The NO3∶PO4 disappearance ratio in waters dominated by P. antarctica (19.0±0.61) was significantly greater than in waters where diatoms were most common (9.52±0.33), and both were significantly different from the Redfield N∶P ratio of 16. Vertical profiles of TDIC suggest that P. antarctica took up 110% more CO2 per mole of PO4 removed than did diatoms, an important consideration for climate models that estimate C uptake from the removal of PO4.

In March 1998 we measured iron in the upper water column and conducted iron- and nutrient-enrichment bottle-incubation experiments in the open-ocean Subantarctic region southwest of Tasmania, Australia. In the Subtropical Convergence Zone (∼42°S, 142°E), silicic acid concentrations were low (< 1.5 µM) in the upper water column, whereas pronounced vertical gradients in dissolved iron concentration (0.12-0.84 nM) were observed, presumably reflecting the interleaving of Subtropical and Subantarctic waters, and mineral aerosol input. Results of a bottle-incubation experiment performed at this location indicate that phytoplankton growth rates were limited by iron deficiency within the iron-poor layer of the euphotic zone. In the Subantarctic water mass (∼46.8°S, 142°E), low concentrations of dissolved iron (0.05-0.11 nM) and silicic acid (< 1 µM) were measured throughout the upper water column, and our experimental results indicate that algal growth was limited by iron deficiency. These observations suggest that availability of dissolved iron is a primary factor limiting phytoplankton growth over much of the Subantarctic Southern Ocean in the late summer and autumn.

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′S, 170°40′W), 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 by a significant drawdown in macronutrients and an approximate 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 bioavailable 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 bioavailable iron from melting sea ice is an important factor regulating phytoplankton production, particularly ice-edge blooms, in seasonally ice-covered Antarctic waters.

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 μg L−1, 0.54 mg L−1 and 2.63 g C m−2 d−1, respectively. Comparable mean concentrations and rates for February were 1.1 μg L−1, 0.29 mg L−1, and 0.78 g C m−2 d−1 (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−2 d−1 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−1 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.

The distribution of phytoplankton biomass and pigments was determined in the Ross Sea during late austral summer 1992. Large gradients in biomass were noted both in the east-west and north-south direction, with maximum particulate matter concentrations occurring in the southwest portion of the Ross Sea. Two xanthophyll pigments dominated the profiles, fucoxanthin (indicative of diatoms) and 19′ hexanoyloxyfucoxanthin (hex) (indicative of the prymnesiophyte Phaeocystis antarctica). Fucoxanthin concentrations were greatest near the coast of Victoria Land and within the northern transect, whereas hex concentrations were highest offshore in the SE Ross Sea. Particulate carbon:chlorophyll a (POC:chl) ratios in diatom-dominated waters of the western Ross Sea were relatively high (210). The POC:chl ratio for Phaeocystis antarctica populations from the SE Ross Sea was substantially less (92) than the ratios observed in diatom-dominated waters. Pigment 14C labeling indicated that phytoplankton carbon in the diatom-dominated northern Ross Sea accounted for approximately 30% of the POC, with a phytoplankton C:chl ratio of 130. Short-term (24 hour) vertical fluxes of pigments ranged from 3 to 40 μg chl a m−2 d−1. Diatom-dominated regions had greater fluxes of phaeophorbides, suggesting that metazoans were the most important grazers at these locations. In contrast, the phaeophytin/total phaeopigment ratio was highest in waters dominated by Phaeocystis antarctica. The distribution of phytoplankton biomass and pigments revealed a spatially variable distribution of taxa, one which clearly has important consequences to food-web dynamics, biogeochemical cycles, and vertical flux patterns in the Ross Sea.

[1] The Ross Sea polynya is among the most productive regions in the Southern Ocean and may constitute a significant oceanic CO2 sink. Based on results from several field studies, this region has been considered seasonally iron limited, whereby a “winter reserve” of dissolved iron (dFe) is progressively depleted during the growing season to low concentrations (∼0.1 nM) that limit phytoplankton growth in the austral summer (December–February). Here we report new iron data for the Ross Sea polynya during austral summer 2005–2006 (27 December–22 January) and the following austral spring 2006 (16 November–3 December). The summer 2005–2006 data show generally low dFe concentrations in polynya surface waters (0.10 ± 0.05 nM in upper 40 m, n = 175), consistent with previous observations. Surprisingly, our spring 2006 data reveal similar low surface dFe concentrations in the polynya (0.06 ± 0.04 nM in upper 40 m, n = 69), in association with relatively high rates of primary production (∼170–260 mmol C m−2 d−1). These results indicate that the winter reserve dFe may be consumed relatively early in the growing season, such that polynya surface waters can become “iron limited” as early as November; i.e., the seasonal depletion of dFe is not necessarily gradual. Satellite observations reveal significant biomass accumulation in the polynya during summer 2006–2007, implying significant sources of “new” dFe to surface waters during this period. Possible sources of this new dFe include episodic vertical exchange, lateral advection, aerosol input, and reductive dissolution of particulate iron.

[1] We calculate net community production (NCP) during summer 2005–2006 and spring 2006 in the Ross Sea using multiple approaches to determine the magnitude and consistency of rates. Water column carbon and nutrient inventories and surface ocean O2/Ar data are compared to satellite-derived primary productivity (PP) estimates and 14C uptake experiments. In spring, NCP was related to stratification proximal to upper ocean fronts. In summer, the most intense C drawdown was in shallow mixed layers affected by ice melt; depth-integrated C drawdown, however, increased with mixing depth. ΔO2/Ar-based methods, relying on gas exchange reconstructions, underestimate NCP due to seasonal variations in surface ΔO2/Ar and NCP rates. Mixed layer ΔO2/Ar requires approximately 60 days to reach steady state, starting from early spring. Additionally, cold temperatures prolong the sensitivity of gas exchange reconstructions to past NCP variability. Complex vertical structure, in addition to the seasonal cycle, affects interpretations of surface-based observations, including those made from satellites. During both spring and summer, substantial fractions of NCP were below the mixed layer. Satellite-derived estimates tended to overestimate PP relative to 14C-based estimates, most severely in locations of stronger upper water column stratification. Biases notwithstanding, NCP-PP comparisons indicated that community respiration was of similar magnitude to NCP. We observed that a substantial portion of NCP remained as suspended particulate matter in the upper water column, demonstrating a lag between production and export. Resolving the dynamic physical processes that structure variance in NCP and its fate will enhance the understanding of the carbon cycling in highly productive Antarctic environments.

As part of the Asian Dust Inputs to the Ocean System (ADIOS) project, atmospheric dust fluxes and primary productivity were monitored during the dusty season (spring) of 1986 at 26°N, 155°W, in the North Pacific Ocean. The arrival of major pulses of dust from Asia was followed by major increases in primary production. Extensive chemical analyses of the atmospheric particles showed that they were iron-rich (10-15%) and, further, that if only a small proportion (e.g. 10%) of this iron dissolved in the euphotic zone, it would be sufficient to support the increases in carbon production at this location. The systematic increases in production noted with increasing depth and time may result from a continual release of iron from the settling particles in the euphotic zone. At all depths, systematic decreases in production followed the initial surge in production, indicating that the phytoplankton may have evolved from being iron-limited to being nitrogen-limited. Comparison of particle concentrations calculated by a particle settling model with primary productivity profiles indicated that mineral particles with settling velocities equivalent to those of 14 to 18-μm-diameter spherical quartz particles were the most likely source for the iron stimulating the increases in primary production.

behaved as a micronutrient with correlations with major nutrients (nitrate, phosphate; r 2 = 0.90, 0.96) until depleted to � 50 pM of strongly complexed cobalt. Co:P utilization ratios were an order of magnitude higher than in the North Pacific, comparable to utilization rates of zinc in other oceanic regions. Cobalt speciation measurements showed that available cobalt decreased over 4 orders of magnitude in this region, with shifts in phytoplankton assemblages occurring at transitions between labile and nonlabile cobalt. Only small changes in total dissolved nickel were observed, and nickel was present in a labile chemical form throughout the region. In the Peru upwelling region, cobalt uptake was highest at the surface and decreased with depth, suggesting phytoplankton uptake was a more important removal mechanism than co-oxidation with microbial manganese oxidation. These findings show the importance of cobalt as a micronutrient and that cobalt scarcity and speciation may be important in influencing phytoplankton species composition in this economically important environment. INDEX TERMS: 1030 Geochemistry: Geochemical cycles (0330); 1050 Geochemistry: Marine geochemistry (4835, 4850); 1065 Geochemistry: Trace elements (3670); 4279 Oceanography: General: Upwelling and convergences