Phytoplankton iron limitation in the Atlantic Southern Ocean driven by seasonal mixed‐layer dynamics

Chlorophyll a and nutrient concentrations along with temperature and salinity values were measured at 22 CTD stations along a 735-km transect running to the northwest of the island of South Georgia, Southern Ocean. Measurements were repeated during five summer surveys (January and February 1994, January 1996, December 1996, January 1998) and one spring survey (October 1997). The transect sampled Sub-Antarctic Zone water in the north, Polar Frontal Zone water and Antarctic Zone water in the south. Chlorophyll a concentrations were lowest to the north of the transect and frequently high (up to 17 mg m−3) in the deep open ocean of the Antarctic Zone. Sub-surface peaks were measured in all zones and chlorophyll a was detectable to a depth of 150 m. There was a clear latitudinal temperature gradient in the near-surface waters (0–50 m), the warmest water occurring in the north (∼12 °C), and the coolest in the Antarctic Zone (∼2 °C). There was also a well-defined latitudinal gradient in summer near-surface silicate concentrations (∼2, 4, and 10 mmol m−3 in the Sub-Antarctic Zone, the Polar Frontal Zone and the Antarctic Zone, respectively), increasing to >20 mmol m−3 near South Georgia. Distinct differences in silicate concentrations were also evident in all three zones to a depth of 500 m. Near-surface nitrate and phosphate concentrations were relatively low to the north of the transect (∼14 and 1 mmol m−3, respectively) and higher in the Polar Frontal Zone and Antarctic Zone (∼18 and 1.4 mmol m−3, respectively). Ammonium and nitrite were restricted to the upper 200 m of the water column, and exhibited sub-surface concentration peaks, the lowest being in the Sub-Antarctic Zone (0.68 and 0.25 mmol m−3, respectively) and the highest in the Antarctic Zone (1.72 and 0.29 mmol m−3, respectively). Surface (∼6 m) spring nutrient measurements provided an indication of pre-bloom conditions; ammonium and nitrite concentrations were low (∼0.27 and 0.28 mmol m−3, respectively), while silicate, nitrate and phosphate concentrations were high and similar to previously measured winter values (e.g. ∼26, 23, 2 mmol m−3, respectively in the Antarctic Zone). Although the values measured were very variable, and there was some evidence of a seasonal growth progression, the chlorophyll a and nutrient distribution patterns were dominated by intercruise (interannual) factors. Approximate nutrient depletions (spring minus summer) appeared similar in the Polar Frontal Zone and Antarctic Zone for nitrate and phosphate, while silicate showed a marked latitudinal increase from north to south throughout the transect. Highest chlorophyll a concentrations coincided with the highest apparent silicate depletions over the deep ocean of the Antarctic Zone. In this area, relatively warm, easterly flowing Antarctic Circumpolar Current water meets cooler, westerly flowing water that is influenced by the Weddell-Scotia Confluence and is rich in nutrients, especially silicate.

Chlorophyll fluorescence, primarily used to derive phytoplankton biomass, has long been an underutilized source of information on phytoplankton physiology. Diel fluctuations in chlorophyll fluorescence are affected by both photosynthetic efficiency and non-photochemical quenching (NPQ), where NPQ is a decrease in fluorescence through the dissipation of excess energy as heat. NPQ variability is linked to iron and light availability, and has the potential to provide important diagnostic information on phytoplankton physiology. Here we establish a relationship between NPQsv (Stern-Volmer NPQ) and indices of iron limitation from nutrient addition experiments in the sub-Antarctic zone (SAZ) of the Atlantic Southern Ocean, through the derivation of NPQmax (the maximum NPQsv value) and αNPQ (the light limited slope of NPQsv). Significant differences were found for both Fv/Fm and αNPQ for iron versus control treatments, with no significant differences for NPQmax. Similar results from CTDs indicated that changes in NPQ were driven by increasing light availability from late July to December, but by both iron and light from January to February. We propose here that variability in αNPQ, which has removed the effect of light availability, can potentially be used as a proxy for iron limitation (as shown here for the Atlantic SAZ), with higher values being associated with greater iron stress. This approach was transferred to data from a buoyancy glider deployment at the same location by utilising the degree of fluorescence quenching as a proxy for NPQGlider, which was plotted against in situ light to determine αNPQ. Seasonal increases in αNPQ are consistent with increased light availability, shoaling of the mixed layer depth (MLD) and anticipated seasonal iron limitation. The transition from winter to summer, when positive net heat flux dominates stratification, was coincident with a 24% increase in αNPQ variability and a switch in the dominant driver from incident PAR to MLD. The dominant scales of αNPQ variability are consistent with fine scale variability in MLD and a significant positive relationship was observed between these two at a ~10 day window. The results emphasise the important role of fine scale dynamics in driving iron supply, particularly in summer when this micronutrient is limiting.

Observations of phytoplankton iron limitation in the world's oceans have primarily been confined to high-nutrient, low-chlorophyll (HNLC) regimes, found in the western equatorial and sub- arctic Pacific, Southern Ocean, and coastal upwelling zones off California and Peru. We investigated the potential for phytoplankton iron limitation in coastal transition zones (50 to 200 km offshore) of the southern California Current System, a weak upwelling regime that is relatively low in nutrients (< 4 µmol nitrate l -1 ) and low in chlorophyll (<1 µg chl a l -1 ). In grow-out incubation experiments con- ducted during summer, July 2003 and 2004, phytoplankton responded to nanomolar iron additions, despite the non-HNLC initial conditions. Observed changes in phytoplankton and nutrient para- meters upon iron addition were significant, although markedly lower in amplitude relative to typical grow-out experiments in HNLC regimes. While we cannot disprove alternate explanations for the observed limitation of phytoplankton growth, such as a proximate grazing control, our results indicate that phytoplankton growth in the southern California Current System is, at times, limited by the supply of iron. Based on our findings and the results of previous studies in this region, we suggest that phytoplankton biomass is generally limited by the supply of nitrate, while iron, directly or indirectly, influences macronutrient utilization, community species composition, and phytoplankton spatial and temporal distribution.

Ecological and biogeographic relationships of class Flavobacteria in the Southern Ocean

Physical controls on biogeochemical zonation in the Southern Ocean

Simulation of chlorophyll and iron supplies in the Sub Antarctic Zone South of Australia

During four summer seasons mesozooplankton community composition and structure in relation to water mass distribution were investigated along a 735-km transect running across the Antarctic Circumpolar Current (ACC) to the north-west of the island of South Georgia, Southern Ocean. Samples were obtained each year during December–January from the top 200 m of the water column at 22 stations spaced 35 km apart. Cluster analysis revealed four station groupings that were geographically consistent with the different water masses identified on the basis of temperature and salinity properties along the transect. A Sub-Antarctic Zone (SAZ) community characterised by low overall plankton abundance was present at the northernmost end of the transect on three of the four cruises, separated from a Polar Frontal Zone (PFZ) community by the Sub-Antarctic Front (SAF). The PFZ community lay between the SAF and the Polar Front (PF) and was characterised by highest overall abundance and little interannual variability. Two Antarctic Zone (AAZ) communities were found south of the PF that, although taxonomically similar, differed in overall abundance. Although there were significant differences in nutrients (e.g. silicate) and phytoplankton (Chl a) between the different water masses, these factors only weakly correlated with plankton community structure. Copepods were the largest contributors to total abundance within all station groupings (median percentage 83–90% of total) and pteropods were also proportionately abundant in the PFZ (11%). With the exception of pteropods (≤11%) and ostracods (≤3%) all other non-copepod taxa contributed <1% to total abundance. All station groups were characterised by varying proportions of a relatively small subset of species, many of which were present throughout the transect, for example, Oithona similis, Ctenocalanus spp., Euchaeta antarctica, and Rhincalanus gigas. Others were particularly characteristic of different station groups, thus Neocalanus tonsus, Clausocalanus ingens, and Calocalanus spp. were characteristic of the SAZ with few in the PFZ and none in the AAZ. Microcalanus pygmaeus was particularly abundant in the AAZ compared to other regions, as was the polychaete, Pelagobia longicirrata. Other taxa, although widespread, tended to be typical of particular water masses, for example, Calanus simillimus and Limacina helicina in the PFZ, Calanoides acutus, appendicularians and P. longicirrata in the AAZ, and ostracods and chaetognaths in the SAZ. The close physical and biological coupling observed across the ACC confirms the frontal zones and particularly the PF as features across which community properties change in the Atlantic sector of the Southern Ocean.

We present a data‐assimilated model of the ocean's phosphorus cycle that is constrained by climatological phosphate, temperature, salinity, sea‐surface height, surface heat and freshwater fluxes, as well as chlorofluorocarbon‐11(CFC‐11) and natural Δ14C. Export production is estimated to be 5.8±2.0×1012 mol P/yr of which (26±6)% originates in the Southern Ocean (SO) south of 40°S. The biological pump efficiency, defined as the proportion of the ocean's phosphate inventory that is regenerated, is (39±7)%. Dividing the SO south of 40°S into a sub‐Antarctic zone (SANTZ) and an Antarctic zone (ANTZ) separated by the latitude of maximum Ekman divergence, we estimate that the SANTZ and ANTZ account, respectively, for (23±5)% and (3±1)% of global export production, (17±4)% and (3±1)% of the regenerated nutrient inventory, and (31±1)% and (43±5)% of the preformed nutrient inventory. Idealized SO nutrient depletion experiments reveal a large‐scale transfer of nutrients into circumpolar and deep waters and from the preformed to the regenerated pool. In accord with the concept of the biogeochemical divide, we find that nutrient drawdown in the ANTZ is more effective than in the SANTZ for increasing the efficiency of the biological pump, while having a smaller impact on production in regions north of 40°S. Complete SO nutrient drawdown would allow the biological pump to operate at 94% efficiency by short circuiting the transport of nutrients in northward Ekman currents, leading to a trapping of nutrients in circumpolar and deep waters that would decrease production outside the SO by approximately 44% while increasing it in the SO by more than 725%.

Phytoplankton growth in the Australian sector of the Southern Ocean, examined by optimising ecosystem model parameters

This study reports detailed silicoflagellate assemblage composition and annual seasonal flux from sediment traps at four locations along a transect across the Southern Ocean frontal systems. The four traps sampled the central Subantarctic Zone (SAZ, 47°S site), the Subantarctic Front (SAF, 51°S site), the Polar Frontal Zone (54°S site) and the Antarctic Zone (61°S site) across the 140°E longitude. Annual silicoflagellate fluxes to the deep ocean exhibited a similar latitudinal trend to those of diatom fluxes reported in previous work, with maxima in the Antarctic Zone and minima in the Subantarctic Zone. The data suggest that, along with diatoms, silicoflagellates are important contributors to biogenic silica export at all sites, particularly in the Subantarctic Zone. Two main silicoflagellate genera were observed, with Stephanocha sp. (previously known as Distephanus) dominating polar waters and Dictyocha sp. important in sub-polar waters. This is consistent with previous use of the Dictyocha / Stephanocha ratio to infer paleotemperatures and monitor shifts in the position of the Polar Frontal Zone in the sedimentary record. It appears possible to further refine the application of this approach by using the ratio between two Dictyocha species, because Dictyocha aculeata dominated at the Subantarctic Front, while Dictyocha stapedia dominated in the central Subantarctic Front. Given the well-defined environmental affinities of both species, a new SAF silicoflagellate index (SAF-SI) based on this ratio is proposed as a useful diagnostic for SAF and SAZ water mass signatures in the Plio-Pleistocene and Holocene sedimentary record.

Spatial and temporal distribution of Fe, Ni, Cu and Pb along 140°E in the Southern Ocean during austral summer 2001/02

The organic carbon particle export to the interior layers in the Southern Ocean in the New Zealand–Tasmania Sector was approximately 170 mmolC m−2 yr−1. The export of particulate inorganic carbon in CaCO3 was 110 mmolC m−2 yr−1 and was contributed mostly by pteropods shells in the Antarctic Zones. The Si flux from biogenic opal at the sub-Antarctic Zone was 67 mmolSi m−2 yr−1 and rapidly increased to the south up to nearly 1 molSi m−2 yr−1 in the Antarctic Zone. The Antarctic Polar Front clearly demarcated the area where the biological pump was driven by CaCO3 to the north and biogenic SiO2 particle export to the south. Summer stratification caused by the sub-zero winter water layer in the Seasonal Ice Zone (SIZ) curtails the zooplankton community and hinders the replenishment of Fe. This hypothesis explains the large organic carbon export with large f- and export ratios at the SIZ and extremely large opal production at the Antarctic Circumpolar Zone. Estimated regeneration rate of CO2 from the export production and settling particulate fluxes of organic carbon in the water column between 100 m to 1 km was about 13 mmolC m−2 d−1 in the Antarctic Zone and Polar Frontal Zone.

Using observational data collected south of Tasmania during 14 austral summer cruises during 1993–2011, we examined the response of sea surface fugacity of carbon dioxide (fCO2) to the Southern Annular Mode (SAM) shift, which occurred around 2000. In the southern part of the Southern Ocean (SO) or the Polar Zone (PZ) and the Polar Frontal Zone (PFZ), fCO2 increased faster at the sea surface than in the atmosphere before the SAM shift, but not after the shift. In the northern part of the SO or the Subantarctic Zone (SAZ), however, surface fCO2 increased faster than atmospheric fCO2 both before and after the shift. The SAM shift had an important influence on the surface fCO2 trend in the PZ and PFZ but not in the SAZ, which we attribute to differences in regional oceanographic processes (upwelling versus nonupwelling). The SAM shift may have reversed the negative trend of SO CO2 uptake.

Spatial and temporal variation and distribution of mesozooplankton in the Drake Passage sampled with the continuous Plankton Recorder

Horizontal and vertical distribution pattern of living radiolarians along a transect from the Southern Ocean to the South Atlantic subtropical region