Regulation of Phytoplankton Carbon to Chlorophyll Ratio By Light, Nutrients and Temperature in the Equatorial Pacific Ocean: A Basin-Scale Model

The complex effects of light, nutrients and temperature lead to a variable carbon to chlorophyll (C:Chl) ratio in phytoplankton cells. Using field data collected in the Equatorial Pacific, we derived a new dynamic model with a non-steady C:Chl ratio as a function of irradiance, nitrate, iron, and temperature. The dynamic model is implemented into a basin-scale ocean circulation-biogeochemistry model and tested in the Equatorial Pacific Ocean. The model reproduces well the general features of phytoplankton dynamics in this region. For instance, the simulated deep chlorophyll maximum (DCM) is much deeper in the western warm pool (similar to 100 m) than in the Eastern Equatorial Pacific (similar to 50 m). The model also shows the ability to reproduce chlorophyll, including not only the zonal, meridional and vertical variations, but also the interannual variability. This modeling study demonstrates that combination of nitrate and iron regulates the spatial and temporal variations in the phytoplankton C:Chl ratio in the Equatorial Pacific. Sensitivity simulations suggest that nitrate is mainly responsible for the high C:Chl ratio in the western warm pool while iron is responsible for the frontal features in the C:Chl ratio between the warm pool and the upwelling region. In addition, iron plays a dominant role in regulating the spatial and temporal variations of the C:Chl ratio in the Central and Eastern Equatorial Pacific. While temperature has a relatively small effect on the C:Chl ratio, light is primarily responsible for the vertical decrease of phytoplankton C:Chl ratio in the euphotic zone.

Modeling Responses of Diatom Productivity and Biogenic Silica Export to Iron Enrichment in the Equatorial Pacific Ocean

Using a three-dimensional physical-biogeochemical model, we have investigated the modeled responses of diatom productivity and biogenic silica export to iron enrichment in the equatorial Pacific, and compared the model simulation with in situ (IronEx II) iron fertilization results. In the eastern equatorial Pacific, an area of 540,000 km(2) was enhanced with iron by changing the photosynthetic efficiency and silicate and nitrogen uptake kinetics of phytoplankton in the model for a period of 20 days. The vertically integrated Chl a and primary production increased by about threefold 5 days after the start of the experiment, similar to that observed in the IronEx II experiment. Diatoms contribute to the initial increase of the total phytoplankton biomass, but decrease sharply after 10 days because of mesozooplankton grazing. The modeled surface nutrients (silicate and nitrate) and TCO(2) anomaly fields, obtained from the difference between the "iron addition'' and "ambient'' (without iron) concentrations, also agreed well with the IronEx II observations. The enriched patch is tracked with an inert tracer similar to the SF6 used in the IronEx II. The modeled depth-time distribution of sinking biogenic silica (BSi) indicates that it would take more than 30 days after iron injection to detect any significant BSi export out of the euphotic zone. Sensitivity studies were performed to establish the importance of fertilized patch size, duration of fertilization, and the role of mesozooplankton grazing. A larger size of the iron patch tends to produce a broader extent and longer-lasting phytoplankton blooms. Longer duration prolongs phytoplankton growth, but higher zooplankton grazing pressure prevents significant phytoplankton biomass accumulation. With the same treatment of iron fertilization in the model, lowering mesozooplankton grazing rate generates much stronger diatom bloom, but it is terminated by Si(OH)(4) limitation after the initial rapid increase. Increasing mesozooplankton grazing rate, the diatom increase due to iron addition stays at minimum level, but small phytoplankton tend to increase. The numerical model experiments demonstrate the value of ecosystem modeling for evaluating the detailed interaction between biogeochemical cycle and iron fertilization in the equatorial Pacific.

Influence of Equatorial Diatom Processes on Si Deposition and Atmospheric CO(2) Cycles at Glacial/Interglacial Timescales

The causes of the glacial cycle remain unknown, although the primary driver is changes in atmospheric CO(2), likely controlled by the biological pump and biogeochemical cycles. The two most important regions of the ocean for exchange of CO(2) with the atmosphere are the equatorial Pacific and the Southern Ocean ( SO), the former a net source and the latter a net sink under present conditions. The equatorial Pacific has been shown to be a Si(OH)(4)-limited ecosystem, a consequence of the low source Si(OH)(4) concentrations in upwelled water that has its origin in the SO. This teleconnection for nutrients between the two regions suggests an oscillatory relationship that may influence or control glacial cycles. Opal mass accumulation rate (MAR) data and delta(15)N measurements in equatorial cores are interpreted with predictions from a one- dimensional Si(OH)(4)-limited ecosystem model (CoSINE) for the equatorial Pacific. The results suggest that equatorial Pacific surface CO(2) processes are in opposite phase to that of the global atmosphere, providing a negative feedback to the glacial cycle. This negative feedback is implemented through the effect of the SO on the equatorial Si(OH)(4) supply. An alternative hypothesis, that the whole ocean becomes Si(OH)(4) poor during cooling periods, is suggested by low opal MAR in cores from both equatorial and Antarctic regions, perhaps as a result of low river input. terminations in this scenario would result from blooms of coccolithophorids triggered by low Si(OH)(4) concentrations.

Size-Fractionated Nitrogen Uptake Measurements in the Equatorial Pacific and Confirmation of the Low Si-High-Nitrate Low-Chlorophyll Condition

The equatorial Pacific Ocean is the largest natural source of CO(2) to the atmosphere, and it significantly impacts the global carbon cycle. Much of the large flux of upwelled CO(2) to the atmosphere is due to incomplete use of the available nitrate (NO(3)) and low net productivity. This high-nutrient low-chlorophyll (HNLC) condition of the equatorial upwelling zone (EUZ) has been interpreted from modeling efforts to be due to low levels of silicate ( Si( OH) 4) that limit the new production of diatoms. These ideas were incorporated into an ecosystem model, CoSINE. This model predicted production by the larger phytoplankton and the picoplankton and effects on air-sea CO(2) fluxes in the Pacific Ocean. However, there were no size-fractionated rates available for verification. Here we report the first size-fractionated new and regenerated production rates (obtained with (15)N - NO(3) and (15)N - NH(4) incubations) for the EUZ with the objective of validating the conceptual basis and functioning of the CoSINE model. Specifically, the larger phytoplankton ( with cell diameters > 5 mu m) had greater rates of new production and higher f-ratios (i.e., the proportion of NO(3) to the sum of NO(3) and NH(4) uptake) than the picoplankton that had high rates of NH(4) uptake and low f-ratios. The way that the larger primary producers are regulated in the EUZ is discussed using a continuous chemostat approach. This combines control of Si(OH)(4) production by supply rate (bottom-up) and control of growth rate ( or dilution) by grazing ( top-down control).

Physical and Biological Controls on the Latitudinal Asymmetry of Surface Nutrients and pCO(2) in the Central and Eastern Equatorial Pacific

Surface nutrients and dissolved inorganic carbon (DIC) in the central (CEP) and eastern equatorial Pacific (EEP) show much higher concentrations to the south than to the north of the equator. In this study, the physical and biological controls on this asymmetry are investigated using a coupled physical-biogeochemical model. Two numerical experiments are conducted to examine the effects of asymmetrical photosynthetic efficiency (a) due to asymmetrical iron supply about the equator. The experiment with asymmetrical photosynthesis produces improved results as compared with historical observations. A nitrate budget analysis suggests that in the EEP the divergence of upwelling waters controls the surface nitrate asymmetry with additional contribution from the South Equatorial Current (SEC) carrying nutrient-rich Peru upwelling water. The changes of a affect the surface nitrate distribution but not the overall asymmetry. The SEC further carries excess nitrate to the west and thus extends the asymmetry in the east to the CEP. In the CEP, however, stronger northward than southward transport tends to reduce the nitrate asymmetry, while the asymmetrical photosynthesis would help to maintain it. Similar processes also control the distributions of surface silicate and DIC in the equatorial Pacific, which is also affected by the air-sea CO(2) exchange. The asymmetrical photosynthesis influences the distribution of surface DIC, pCO(2), and the air-sea CO(2) flux, by redistributing about 20% CO(2) flux from the north to the south of the equator. Owing to the adjustment of air-sea CO(2) flux, however, the net surface DIC change is smaller than the direct change associated with primary production.