Iron-limited Waters Research Articles

On the Variability of Equatorial Pacific Nitrate and Iron Utilization

The tropical Pacific is one of the largest ocean regions on Earth where the trace element iron limits new primary production and therefore the efficiency of carbon export to the deep sea. Although there is a long history of marine biogeochemical research in the tropical Pacific, recent advancements using GEOTRACES key parameters such as iron and nitrate isotopes (nitrate δ15N and δ18O) make this a good time to review the current understanding of tropical Pacific nitrate dynamics—how both regional subsurface nitrate characteristics and surface ocean nitrate utilization change with time. While this article provides a comprehensive overview of the biological, chemical, and physical processes shaping equatorial Pacific subsurface-to-surface nutrients, it principally explores the findings from the first nitrate isotope time series in iron-limited high nutrient, low chlorophyll waters. Results indicate that the preferential recycling of bioavailable iron within the euphotic zone is required to explain even the lowest observed nitrate utilization in the eastern equatorial Pacific (EEP). Furthermore, because seasonal-to-interannual nitrate utilization variability in the EEP cannot be driven by changes in iron supply, this work argues that iron recycling (and therefore bioavailable iron) is modulated by upwelling rate changes, creating a predicted and recently observed spectrum of iron limitation in the iron-limited EEP surface waters. In other words, upper ocean physics overwhelmingly dominates seasonal-to-interannual nitrate utilization in the iron-​limited EEP. This new understanding of nitrate utilization in iron-limited waters helps to explain long-term changes in past equatorial Pacific nitrate utilization obtained via sedimentary proxy records and potentially complicates the efficacy of future iron fertilization of the equatorial Pacific.

Comparative Thermophysiology of Marine Synechococcus CRD1 Strains Isolated From Different Thermal Niches in Iron-Depleted Areas.

Marine Synechococcus cyanobacteria are ubiquitous in the ocean, a feature likely related to their extensive genetic diversity. Amongst the major lineages, clades I and IV preferentially thrive in temperate and cold, nutrient-rich waters, whilst clades II and III prefer warm, nitrogen or phosphorus-depleted waters. The existence of such cold (I/IV) and warm (II/III) thermotypes is corroborated by physiological characterization of representative strains. A fifth clade, CRD1, was recently shown to dominate the Synechococcus community in iron-depleted areas of the world ocean and to encompass three distinct ecologically significant taxonomic units (ESTUs CRD1A-C) occupying different thermal niches, suggesting that distinct thermotypes could also occur within this clade. Here, using comparative thermophysiology of strains representative of these three CRD1 ESTUs we show that the CRD1A strain MITS9220 is a warm thermotype, the CRD1B strain BIOS-U3-1 a cold temperate thermotype, and the CRD1C strain BIOS-E4-1 a warm temperate stenotherm. Curiously, the CRD1B thermotype lacks traits and/or genomic features typical of cold thermotypes. In contrast, we found specific physiological traits of the CRD1 strains compared to their clade I, II, III, and IV counterparts, including a lower growth rate and photosystem II maximal quantum yield at most temperatures and a higher turnover rate of the D1 protein. Together, our data suggests that the CRD1 clade prioritizes adaptation to low-iron conditions over temperature adaptation, even though the occurrence of several CRD1 thermotypes likely explains why the CRD1 clade as a whole occupies most iron-limited waters.

Widespread phytoplankton blooms triggered by 2019-2020 Australian wildfires.

Droughts and climate-change-driven warming are leading to more frequent and intense wildfires1-3, arguably contributing to the severe 2019-2020 Australian wildfires4. The environmental and ecological impacts of the fires include loss of habitats and the emission of substantial amounts of atmospheric aerosols5-7.Aerosol emissions from wildfires can lead to the atmospheric transport of macronutrients and bio-essential trace metals such as nitrogen and iron, respectively8-10. It has been suggested that the oceanic deposition of wildfire aerosols can relieve nutrient limitations and, consequently, enhance marine productivity11,12, but direct observations are lacking. Here we use satellite and autonomous biogeochemical Argo float data to evaluate the effect of 2019-2020 Australian wildfire aerosol deposition on phytoplankton productivity. We find anomalously widespread phytoplankton blooms from December 2019 to March 2020 in the Southern Ocean downwind of Australia. Aerosol samples originating from the Australian wildfires contained a high iron content and atmospheric trajectories show that these aerosols were likely to be transported to the bloom regions, suggesting that the blooms resulted from the fertilization of the iron-limited waters of the Southern Ocean. Climate models project more frequent and severe wildfires in many regions1-3. A greater appreciation of the links between wildfires, pyrogenic aerosols13, nutrient cycling and marine photosynthesis could improve our understanding of the contemporary and glacial-interglacial cycling of atmospheric CO2 and the global climate system.

Shunt or shuttle

Nutrient availability influences the susceptibility of marine primary producers to viral infection. For diatoms in iron-limited waters, reduced infection rates impact marine biogeochemical cycles by enhancing the flux of material to depth.

Endocytosis-mediated siderophore uptake as a strategy for Fe acquisition in diatoms.

Phytoplankton growth is limited in vast oceanic regions by the low bioavailability of iron. Iron fertilization often results in diatom blooms, yet the physiological underpinnings for how diatoms survive in chronically iron-limited waters and outcompete other phytoplankton when iron becomes available are unresolved. We show that some diatoms can use siderophore-bound iron, and exhibit a species-specific recognition for siderophore types. In Phaeodactylum tricornutum, hydroxamate siderophores are taken up without previous reduction by a high-affinity mechanism that involves binding to the cell surface followed by endocytosis-mediated uptake and delivery to the chloroplast. The affinity recorded is the highest ever described for an iron transport system in any eukaryotic cell. Collectively, our observations suggest that there are likely a variety of iron uptake mechanisms in diatoms besides the well-established reductive mechanism. We show that iron starvation-induced protein 1 (ISIP1) plays an important role in the uptake of siderophores, and through bioinformatics analyses we deduce that this protein is largely diatom-specific. We quantify expression of ISIP1 in the global ocean by querying the Tara Oceans atlas of eukaryotic genes and show a link between the abundance and distribution of diatom-associated ISIP1 with ocean provinces defined by chronic iron starvation.

Nutrient Cycling: Are Antarctic Krill a Previously Overlooked Source in the Marine Iron Cycle?

Iron limits phytoplankton growth in large areas of the Southern Ocean. A new study shows that Antarctic krill play a crucial role in the recycling of iron in the iron-limited waters.

A preliminary model of iron fertilisation by baleen whales and Antarctic krill in the Southern Ocean: Sensitivity of primary productivity estimates to parameter uncertainty

Large marine animals may play a crucial role in storing and recycling bioavailable iron in surface waters by consuming iron-rich prey and subsequent defecation of iron that is excess to their requirements. This biological recycling of iron could enhance primary productivity in iron-limited waters. However, quantifying the effects of marine animals on ocean primary productivity remains challenging because of a limited understanding of the key biogeochemical processes involved. In this paper, we develop a preliminary model that explores these uncertainties and examines the potential effects of historical populations of blue, fin and humpback whales, and the biomass of Antarctic krill required to support the whale populations, on primary productivity in the Southern Ocean.Our results suggest that, despite conservative estimates for key processes in our model, pre-exploitation populations of blue whales and, to a lesser extent fin and humpback whales, could have contributed to iron recycling, resulting in enhanced phytoplankton production in iron-limited Southern Ocean waters. Iron-rich defecation by un-exploited whale populations in the Southern Ocean, and the biomass Antarctic krill required to support them, could have resulted in a contribution to primary productivity of between 1.5×10−4 to 23.4gCm−2yr−1 (blue whales), 1.4×10−4 to 13.9gCm−2yr−1 (fin whales), and 2.4×10−5 to 1.7gCm−2yr−1 (humpback whales). However, only when all parameter estimates are at their upper limits does there appear to be this significant role for whales in enhancing primary productivity, and thus we need to assess the likelihood of these values arising.The high degree of uncertainty around the magnitude of these increases in primary productivity is mainly due to our limited quantitative understanding of key biogeochemical processes. To reduce uncertainty regarding the effect of whales on Southern Ocean primary productivity, future research will need to refine our understanding of five influential model parameters: iron content in krill; krill consumption rates by whales; persistence of whale faecal iron in the photic zone; bioavailability of this retained iron; and the carbon-to-iron ratio of phytoplankton.

A Novel Protein, Ubiquitous in Marine Phytoplankton, Concentrates Iron at the Cell Surface and Facilitates Uptake

Numerous cellular functions including respiration require iron. Plants and phytoplankton must also maintain the iron-rich photosynthetic electron transport chain, which most likely evolved in the iron-replete reducing environments of the Proterozoic ocean [1]. Iron bioavailability has drastically decreased in the contemporary ocean [1], most likely selecting for the evolution of efficient iron acquisition mechanisms among modern phytoplankton. Mesoscale iron fertilization experiments often result in blooms dominated by diatoms [2], indicating that diatoms have adaptations that allow survival in iron-limited waters and rapid multiplication when iron becomes available. Yet the genetic and molecular bases are unclear, as very few iron uptake genes have been functionally characterized from marine eukaryotic phytoplankton, and large portions of diatom iron starvation transcriptomes are genes encoding unknown functions [3-5]. Here we show that the marine diatom Phaeodactylum tricornutum utilizes ISIP2a to concentrate Fe(III) at the cell surface as part of a novel, copper-independent and thermodynamically controlled iron uptake system. ISIP2a is expressed in response to iron limitation several days prior to the induction of ferrireductase activity, and it facilitates significant Fe(III) uptake during the initial response to Fe limitation. ISIP2a is able to directly bind Fe(III) and increase iron uptake when heterologously expressed, whereas knockdown of ISIP2a in P. tricornutum decreases iron uptake, resulting in impaired growth and chlorosis during iron limitation. ISIP2a is expressed by diverse marine phytoplankton, indicating that it is an ecologically significant adaptation to the unique nutrient composition of marine environments.

The impact of iron limitation on the physiology of the Antarctic diatom Chaetoceros simplex.

Iron availability strongly governs the growth of Southern Ocean phytoplankton. To investigate how iron limitation affects photosynthesis as well as the uptake of carbon and iron in the Antarctic diatom Chaetoceros simplex, a combination of chlorophyll a fluorescence measurements and radiotracer incubations in the presence and absence of chemical inhibitors was conducted. Iron limitation in C. simplex led to a decline in growth rates, photochemical efficiency and structural changes in photosystem II (PSII), including a reorganisation of photosynthetic units in PSII and an increase in size of the functional absorption cross section of PSII. Iron-limited cells further exhibited a reduced plastoquinone pool and decreased photosynthetic electron transport rate, while non-photochemical quenching and relative xanthophyll pigment content were strongly increased, suggesting a photoprotective response. Additionally, iron limitation resulted in a strong decline in carbon fixation and thus the particulate organic carbon quotas. Inhibitor studies demonstrated that, independent of the iron supply, carbon fixation was dependent on internal, but not on extracellular carbonic anhydrase activity. Orthovanadate more strongly inhibited iron uptake in iron-limited cells, indicating that P-type ATPase transporters are involved in iron uptake. The stronger reduction in iron uptake by ascorbate in iron-limited cells suggests that the re-oxidation of iron is required before it can be taken up and further supports the presence of a high-affinity iron transport pathway. The measured changes to photosystem architecture and shifts in carbon and iron uptake strategies in C. simplex as a result of iron limitation provide evidence for a complex interaction of these processes to balance the iron requirements for photosynthesis and carbon demand for sustained growth in iron-limited waters.

Geochemical evidence of oceanic iron fertilization by the Kasatochi volcanic eruption in 2008 and the potential impacts on Pacific sockeye salmon

The Kasatochi volcanic eruption that occurred in the central Aleutian Islands in Alaska, USA, in August 2008 is thought to have induced a massive diatom bloom in the iron-limited waters of the Gulf of Alaska, which potentially affected the oceanic food web by increasing the abundance of zooplankton and sockeye salmon Oncorhynchus nerka in the northeast Pacific Ocean. We report the first seawater experiments involving volcanic ash ejected from the Kasatochi eruption, showing that the ash released 61 to 83 nmol Fe, 374 to 410 nmol NO3-, 5 to 6 nmol PO43- and 170 to 585 nmol SiO2 when it contacted seawater. Our study suggests that the amount of iron released from Kasatochi ash (an increase of 2.0 to 2.8 nM Fe) was indeed sufficient to cause the observed phytoplankton bloom in the northeastern Pacific Gyre, while the impact of macronutrient release was minimal. We further evaluated the multiple, interdependent processes in the oceanic food web related to the diatom bloom, involving the ocean survival of juvenile salmon that entered the northeast Pacific Ocean in the summer of 2008.

Frustule‐related gene transcription and the influence of diatom community composition on silica precipitation in an iron‐limited environment

A microcosm study in iron‐limited waters of the northeast subarctic Pacific Ocean was conducted to examine how iron availability affects the frustule‐related response of individual diatoms and thus the total quantity of silica precipitated by the community. New silica precipitated per cell was estimated using the fluorescent cell stain 2‐(4‐pyridyl)‐5{[4‐dimethylaminoethyl‐aminocarbamoyl)‐methoxy]phenyl}oxazole (PDMPO). Differences in new silica precipitation within a particular genus before and after iron enrichment were small compared to differences among genera, indicating that the quantity of total silica precipitated is particularly sensitive to community composition. Transcriptional patterns of genes encoding silicon transporters, aminopropyltransferases, chitin synthases, and a protein with uncharacterized function were measured in natural populations to identify indicators of the frustule‐related responses of different genera to iron limitation. Transcripts associated with silicon transporters were the most readily detectable in three metatranscriptome datasets and were capable of resolving species composition shifts and physiological responses. Silicon transporter transcripts from a distinct phylogenetic clade were most abundant in the iron‐limited community, and transcripts from a separate clade were more abundant in the community that bloomed after iron enrichment. Transcripts of the gene present in the iron‐limited community were also more abundant in iron‐limited laboratory cultures of Pseudo‐nitzschia multiseries, suggesting that this gene plays a role in silicon uptake during iron limitation. The responses of individual cells, as detected in this study, determine how the community influences silicon cycling in iron‐limited environments.

Regional Variation in Lytic and Lysogenic Viral Infection in the Southern Ocean and Its Contribution to Biogeochemical Cycling

Lytic and lysogenic viral infection was investigated throughout the Southern Ocean at sites spanning the sub-Antarctic zone, the Antarctic Circumpolar Current, and an Antarctic continental sea. Higher lytic virus activity was recorded in the more productive sub-Antarctic zone than in the iron-limited waters of the Antarctic Circumpolar Current during two transects. Reduced lytic viral activity in the Antarctic Circumpolar Current was combined with a shift toward lysogenic infection, probably resulting from the lower concentration of potential prokaryotic hosts. Superimposed on this variation, lytic viral production was lower in a transect completed in the Drake Passage in autumn (1.8 × 10(8) to 1.5 × 10(9) liter(-1) day(-1)) than over the Greenwich Meridian during summer (5.1 × 10(8) to 2.0 × 10(10) cells liter(-1) day(-1)), indicating that viral activity is linked to the overall seasonal fluctuations in biotic activity. Interestingly, while prokaryotic abundance was lowest in the coastal Weddell Sea, levels of bacterial and lytic viral production (4.3 × 10(8) to 1.7 × 10(10) cells liter(-1) day(-1)) in this area were similar to those of the other zones. This may explain the weak relationship between the distribution of prokaryotes and chlorophyll in the Weddell Sea, as a high turnover of prokaryotic biomass may have been stimulated by the availability of substrates in the form of viral lysate. With estimated carbon and iron releases of 0.02 to 7.5 μg liter(-1) day(-1) and 1.5 to 175.7 pg liter(-1) day(-1), respectively, viral activity in the Southern Ocean is shown to be a major contributor to satisfying the elemental requirements of microbes, notably prokaryotes in the Weddell Sea and phytoplankton in the sub-Antarctic zone.

Glacial flour dust storms in the Gulf of Alaska: Hydrologic and meteorological controls and their importance as a source of bioavailable iron

[1] Iron is an essential micronutrient that limits primary productivity in much of the ocean, including the Gulf of Alaska (GoA). However, the processes that transport iron to the ocean surface are poorly quantified. We combine satellite and meteorological data to provide the first description of widespread dust transport from coastal Alaska into the GoA. Dust is frequently transported from glacially-derived sediment at the mouths of several rivers, the most prominent of which is the Copper River. These dust events occur most frequently in autumn, when coastal river levels are low and riverbed sediments are exposed. The dust plumes are transported several hundred kilometers beyond the continental shelf into iron-limited waters. We estimate the mass of dust transported from the Copper River valley during one 2006 dust event to be between 25–80 ktons. Based on conservative estimates, this equates to a soluble iron loading of 30–200 tons. We suggest the soluble Fe flux from dust originating in glaciofluvial sediment deposits from the entire GoA coastline is two to three times larger, and is comparable to the annual Fe flux to GoA surface waters from eddies of coastal origin. Given that glaciers are retreating in the coastal GoA region and in other locations, it is important to examine whether fluxes of dust are increasing from glacierized landscapes to the ocean, and to assess the impact of associated Fe on marine ecosystems.

Iron regeneration and organic iron(III)-binding ligand production during in situ zooplankton grazing experiment

To elucidate iron regeneration and organic iron(III)-binding ligand formation during microzooplankton and copepod grazing on phytoplankton, incubation experiments were conducted in the western subarctic Pacific. During 8 days of dark incubation of ambient water and that amended with plankton concentrate, dissolved iron and organic iron(III)-binding ligands accumulated, approximately proportionally to the decrease in chlorophyll a. The observed increases in dissolved iron concentration were much greater than those expected from the consumption of phytoplankton biomass and previously reported Fe:C value of cultured algal cells, suggesting resolution from colloidal or particulate iron adsorbed onto the algal cell surface. When copepods were added to the ambient water, organic iron(III)-binding ligands accumulated more rapidly than in the control receiving no copepod addition, although consumed phytoplankton biomass was comparable between the two treatments. Bioassay experiment using filtrates collected from the incubation experiment showed that organic ligands formed during microzooplankton grazing reduced the iron bioavailability to phytoplankton and suppressed their growth. Moreover, picoplankton Synechococcus sp. and Micromonas pusilla were more suppressed by the organic ligands than the diatom Thalassiosira weissflogii. In conclusion, through microzooplankton and copepod grazing on phytoplankton, organic iron(III)-binding ligands as well as regenerated iron are released into the ambient seawater. Because the ligands lower iron bioavailability to phytoplankton through complexation and the degree of availability reduction varies among phytoplankton species, grazing by zooplankton can shift phytoplankton community structure in iron-limited waters.

Annual ecosystem variability in the tropical Indian Ocean: Results of a coupled bio-physical ocean general circulation model

Abstract A coupled, 3-D bio-physical ocean general circulation model has been employed to study biogeochemical variability in the Indian Ocean. The model's ecosystem includes two size classes each of phytoplankton, zooplankton and detritus, and allows iron- and nitrogen-limited phytoplankton growth to develop. Comparison of SeaWiFS and modeled chlorophyll a demonstrates that the model successfully captures seasonal inter-regional contrasts in phytoplankton distribution. Model validation was performed against SeaWiFS ocean color, NODC nitrate, and US JGOFS observations from the Arabian Sea. These data were used to objectively evaluate several ecosystem model modifications that improved the solution. Phytoplankton speciation shifts were generally consistent with those observed in the Arabian Sea. The model shows pronounced basinwide spatio-temporal variability in iron availability; surface waters in the western equatorial and southern tropical regions always tend toward iron limitation, while the Bay of Bengal and the eastern Arabian Sea are largely iron replete. The model also indicates that surface waters in the western Arabian Sea are prone to seasonal iron limitation, despite proximity to regional dust sources. Phytoplankton blooms that manifest as a response to the Somali Current show better agreement with the SeaWiFS data than do previous modeling studies and biogeochemical variability associated with eastward propagation of the semiannual Wyrtki Jet is described for the first time. In the southern tropical region, the model indicates that austral winter Ekman pumping and westward propagating Rossby waves are the primary means of nutrient supply to these iron-limited waters. Finally, inter-regional connections associated with the propagation of the Somali Current and the Wyrtki Jet out of their respective source regions are shown to be a primary influence on seasonal biogeochemical variability in the southern Bay of Bengal. The results presented here establish a need for additional studies that apply such a basinwide approach to further advance our understanding of how bioavailable iron distribution, physical–biological interactions and inter-regional connections affect biogeochemical processes in the Indian Ocean.

Iron transport by mesoscale Haida eddies in the Gulf of Alaska

Abstract A two-year study of the iron distributions in anticyclonic mesoscale Haida eddies of the eastern North Pacific Ocean has shown that such eddies are a major source of iron to the high nitrate–low chlorophyll (HNLC) waters of the central Gulf of Alaska basin. These eddies, which are typically about 100 km in diameter, form off the west coast of Canada in winter and then track roughly westward into the open ocean. Therefore, they carry large quantities of iron-rich coastal waters into iron-limited waters of the oligotrophic gyre. When the eddies had first formed, their dissolved iron concentrations were nearly two orders of magnitude higher than is typically observed at Station Papa (50°N, 145°W; 3 n m versus less than 0.05 n m ), and the total iron in the eddies surface waters (10–40 m) was more than 13 n m higher than open ocean waters of the Alaska gyre (14 n m versus 0.5 n m ). While the overall iron content of the eddies decreased rapidly during the first year after they formed, the eddy we were able to track the longest still contained 1.5–2 times more iron than the surrounding waters 16 months after its formation. Therefore, although iron concentrations in the surface mixed layers of the eddies drop to levels observed outside very rapidly (within 4 months), upward transport along isopycnals and upwelling due to eddy decay, as well as vertical advective and diffusive transports typical of North Pacific waters, probably provide steady fluxes of iron into the euphotic zone from the still iron-rich eddy core waters throughout the lifetime of the eddy.

Determination of organic complexation of cobalt in seawater by cathodic stripping voltammetry

A method was developed to determine the extent of cobalt complexation with organic ligands in seawater, and applied to samples from the North Eastern Atlantic Ocean. The cobalt speciation was determined using catalytic cathodic stripping voltammetry with ligand competition against the adsorptive ligand (nioxime). Optimised conditions include a nioxime concentration of 200 nM, 0.07 M ammonia/ammonium chloride pH buffer (pH 9.1), and 0.5 M nitrite. The stability of the mixed complex of cobalt, nioxime and ammonia was calibrated against EDTA. The cobalt speciation in the open-ocean surface waters was found to be dominated by complexation to natural organic ligands with conditional stability constants (log KCoL′) ranging between 15.6 and 16.1 and with ligand concentrations between 22 and 60 pM. The cobalt concentrations varied between 25 pM in the open-ocean waters and 103 pM in the English Channel, and were less than the ligand concentrations in many of the surface oceanic waters. The ligand concentration at depths between 30 and 115 m (average 22 pM) was lower than that of cobalt, whereas at other depths (shallower as well as deeper), the ligand concentrations were between 26 and 31 pM, sometimes greater than the cobalt concentrations. The cobalt and ligand concentrations in these waters are finely balanced, strongly binding all cobalt when the ligands are in excess. Free Co2+ concentrations calculated for the open-ocean surface waters are extremely low (<5 fM) which could cause cobalt limitation to certain phytoplankton such as coccolithophores and cyanobacteria if the organic complexes are unavailable. If this organic complexation of cobalt occurs also in other regions characterised by low cobalt concentrations, iron-limited waters in high-nitrate low-chlorophyll regions may be cobalt-limited too.

Influence of iron availability on nutrient consumption ratio of diatoms in oceanic waters

The major nutrients (nitrate, phosphate and silicate) needed for phytoplankton growth are abundant in the surface waters of the subarctic Pacific, equatorial Pacific and Southern oceans, but this growth is limited by the availability of iron1,2,3,4,5. Under iron-deficient conditions, phytoplankton exhibit reduced uptake of nitrate6 and lower cellular levels of carbon, nitrogen and phosphorus7. Here I describe seawater and culture experiments which show that iron limitation can also affect the ratio of consumed silicate to nitrate and phosphate. In iron-limited waters from all three of the aforementioned environments, addition of iron to phytoplankton assemblages in incubation bottles halved the silicate:nitrate and silicate:phosphate consumption ratios, in spite of the preferential growth of diatoms (silica-shelled phytoplankton). The nutrient consumption ratios of the phytoplankton assemblage from the Southern Ocean were similar to those of an iron-deficient laboratory culture of Antarctic diatoms, which exhibit increased cellular silicon or decreased cellular nitrogen and phosphorus in response to iron limitation. Iron limitation therefore increases the export of biogenic silicon, relative to nitrogen and phosphorus, from the surface to deeper waters. These findings suggest how the sedimentary records of carbon and silicon deposition in the glacial Southern Ocean8 can be consistent with the idea that changes in productivity, and thus in drawdown of atmospheric CO2, during the last glaciation were stimulated by changes in iron inputs from atmospheric dust.

Contents volume 3, 1978/1979

A coupled, 3-D bio-physical ocean general circulation model has been employed to study biogeochemical variability in the Indian Ocean. The model's ecosystem includes two size classes each of phytoplankton, zooplankton and detritus, and allows iron- and nitrogen-limited phytoplankton growth to develop. Comparison of SeaWiFS and modeled chlorophyll a demonstrates that the model successfully captures seasonal inter-regional contrasts in phytoplankton distribution. Model validation was performed against SeaWiFS ocean color, NODC nitrate, and US JGOFS observations from the Arabian Sea. These data were used to objectively evaluate several ecosystem model modifications that improved the solution. Phytoplankton speciation shifts were generally consistent with those observed in the Arabian Sea. The model shows pronounced basinwide spatio-temporal variability in iron availability; surface waters in the western equatorial and southern tropical regions always tend toward iron limitation, while the Bay of Bengal and the eastern Arabian Sea are largely iron replete. The model also indicates that surface waters in the western Arabian Sea are prone to seasonal iron limitation, despite proximity to regional dust sources. Phytoplankton blooms that manifest as a response to the Somali Current show better agreement with the SeaWiFS data than do previous modeling studies and biogeochemical variability associated with eastward propagation of the semiannual Wyrtki Jet is described for the first time. In the southern tropical region, the model indicates that austral winter Ekman pumping and westward propagating Rossby waves are the primary means of nutrient supply to these iron-limited waters. Finally, inter-regional connections associated with the propagation of the Somali Current and the Wyrtki Jet out of their respective source regions are shown to be a primary influence on seasonal biogeochemical variability in the southern Bay of Bengal. The results presented here establish a need for additional studies that apply such a basinwide approach to further advance our understanding of how bioavailable iron distribution, physical–biological interactions and inter-regional connections affect biogeochemical processes in the Indian Ocean.