Michael J. Follows

Emergent biogeography of microbial communities in a model ocean.

A marine ecosystem model seeded with many phytoplankton types, whose physiological traits were randomly assigned from ranges defined by field and laboratory data, generated an emergent community structure and biogeography consistent with observed global phytoplankton distributions. The modeled organisms included types analogous to the marine cyanobacterium Prochlorococcus. Their emergent global distributions and physiological properties simultaneously correspond to observations. This flexible representation of community structure can be used to explore relations between ecosystems, biogeochemical cycles, and climate change.

Oceanic sources, sinks, and transport of atmospheric CO2

We synthesize estimates of the contemporary net air-sea CO2 flux on the basis of an inversion of interior ocean carbon observations using a suite of 10 ocean general circulation models (Mikaloff Fletcher et al., 2006, 2007) and compare them to estimates based on a new climatology of the air-sea difference of the partial pressure of CO2 (pCO2) (Takahashi et al., 2008). These two independent flux estimates reveal a consistent description of the regional distribution of annual mean sources and sinks of atmospheric CO2 for the decade of the 1990s and the early 2000s with differences at the regional level of generally less than 0.1 Pg C a−1. This distribution is characterized by outgassing in the tropics, uptake in midlatitudes, and comparatively small fluxes in thehigh latitudes. Both estimates point toward a small (∼ −0.3 Pg C a−1) contemporary CO2 sink in the Southern Ocean (south of 44°S), a result of the near cancellation between a substantial outgassing of natural CO2 and a strong uptake of anthropogenic CO2. A notable exception in the generally good agreement between the two estimates exists within the Southern Ocean: the ocean inversion suggests a relatively uniform uptake, while the pCO2-based estimate suggests strong uptake in the region between 58°S and 44°S, and a source in the region south of 58°S. Globally and for a nominal period between 1995 and 2000, the contemporary net air-sea flux of CO2 is estimated to be −1.7 ± 0.4 Pg C a−1 (inversion) and −1.4 ± 0.7 Pg C a−1 (pCO2-climatology), respectively, consisting of an outgassing flux of river-derived carbon of ∼+0.5 Pg C a−1, and an uptake flux of anthropogenic carbon of −2.2 ± 0.3 Pg C a−1 (inversion) and −1.9 ± 0.7 Pg C a−1 (pCO2-climatology). The two flux estimates also imply a consistent description of the contemporary meridional transport of carbon with southward ocean transport throughout most of the Atlantic basin, and strong equatorward convergence in the Indo-Pacific basins. Both transport estimates suggest a small hemispheric asymmetry with a southward transport of between −0.2 and −0.3 Pg C a−1 across the equator. While the convergence of these two independent estimates is encouraging and suggests that it is now possible to provide relatively tight constraints for the net air-sea CO2 fluxes at the regional basis, both studies are limited by their lack of consideration of long-term changes in the ocean carbon cycle, such as the recent possible stalling in the expected growth of the Southern Ocean carbon sink.

Evaluation of ocean carbon cycle models with data-based metrics

New radiocarbon and chlorofluorocarbon-11 data from the World Ocean Circulation Experiment are used to assess a suite of 19 ocean carbon cycle models. We use the distributions and inventories of these tracers as quantitative metrics of model skill and find that only about a quarter of the suite is consistent with the new data-based metrics. This should serve as a warning bell to the larger community that not all is well with current generation of ocean carbon cycle models. At the same time, this highlights the danger in simply using the available models to represent the state-of-the-art modeling without considering the credibility of each model.

Inverse estimates of anthropogenic CO2 uptake, transport, and storage by the ocean

deviation of the models weighted by a CFC-based model skill score, which reduces the error range and emphasizes those models that have been shown to reproduce observed tracer concentrations most accurately. The greatest anthropogenic CO2 uptake occurs in the Southern Ocean and in the tropics. The flux estimates imply vigorous northward transport in the Southern Hemisphere, northward cross-equatorial transport, and equatorward transport at high northern latitudes. Compared with forward simulations, we find substantially more uptake in the Southern Ocean, less uptake in the Pacific Ocean, and less global uptake. The large-scale spatial pattern of the estimated flux is generally insensitive to possible biases in the data and the models employed. However, the global uptake scales approximately linearly with changes in the global anthropogenic CO2 inventory. Considerable uncertainties remain in some regions, particularly the Southern Ocean.

Evaluation of ocean model ventilation with CFC-11: comparison of 13 global ocean models

We compared the 13 models participating in the Ocean Carbon Model Intercomparison Project (OCMIP) with regards to their skill in matching observed distributions of CFC-11. This analysis characterizes the abilities of these models to ventilate the ocean on timescales relevant for anthropogenic CO2 uptake. We found a large range in the modeled global inventory (±30%), mainly due to differences in ventilation from the high latitudes. In the Southern Ocean, models differ particularly in the longitudinal distribution of the CFC uptake in the intermediate water, whereas the latitudinal distribution is mainly controlled by the subgrid-scale parameterization. Models with isopycnal diffusion and eddy-induced velocity parameterization produce more realistic intermediate water ventilation. Deep and bottom water ventilation also varies substantially between the models. Models coupled to a sea-ice model systematically provide more realistic AABW formation source region; however these same models also largely overestimate AABW ventilation if no specific parameterization of brine rejection during sea-ice formation is included. In the North Pacific Ocean, all models exhibit a systematic large underestimation of the CFC uptake in the thermocline of the subtropical gyre, while no systematic difference toward the observations is found in the subpolar gyre. In the North Atlantic Ocean, the CFC uptake is globally underestimated in subsurface. In the deep ocean, all but the adjoint model, failed to produce the two recently ventilated branches observed in the North Atlantic Deep Water (NADW). Furthermore, simulated transport in the Deep Western Boundary Current (DWBC) is too sluggish in all but the isopycnal model, where it is too rapid.

Single-cell genomics reveals hundreds of coexisting subpopulations in wild Prochlorococcus.

Cyanobacterial Diversity What does it mean to be a global species? The marine cyanobacterium Prochlorococcus is ubiquitous and, arguably, the most abundant and productive of all living organisms. Although to our eyes the seas look uniform, to a bacterium the oceans bulk is a plethora of microhabitats, and by large-scale single-cell genomic analysis of uncultured cells, Kashtan et al. (p. 416; see the Perspective by Bowler and Scanlan) reveal that Prochlorococcus has diversified to match. This “species” constitutes a mass of subpopulations—each with million-year ancestry—that vary seasonally in abundance. The subpopulations in turn have clades nested within that show covariation between sets of core alleles and variable gene content, indicating flexibility of responses to rapid environmental changes. Large sets of coexisting populations could be a general feature of other free-living bacterial species living in highly mixed habitats. Covariation between the core alleles and flexible gene content of a marine cyanobacterium underpins vast diversity. [Also see Perspective by Bowler and Scanlan] Extensive genomic diversity within coexisting members of a microbial species has been revealed through selected cultured isolates and metagenomic assemblies. Yet, the cell-by-cell genomic composition of wild uncultured populations of co-occurring cells is largely unknown. In this work, we applied large-scale single-cell genomics to study populations of the globally abundant marine cyanobacterium Prochlorococcus. We show that they are composed of hundreds of subpopulations with distinct “genomic backbones,” each backbone consisting of a different set of core gene alleles linked to a small distinctive set of flexible genes. These subpopulations are estimated to have diverged at least a few million years ago, suggesting ancient, stable niche partitioning. Such a large set of coexisting subpopulations may be a general feature of free-living bacterial species with huge populations in highly mixed habitats.

Evaluating global ocean carbon models: The importance of realistic physics

A suite of standard ocean hydrographic and circulation metrics are applied to the equilibrium physical solutions from 13 global carbon models participating in phase 2 of the Ocean Carbon-cycle Model Intercomparison Project (OCMIP-2). Model-data comparisons are presented for sea surface temperature and salinity, seasonal mixed layer depth, meridional heat and freshwater transport, 3-D hydrographic fields, and meridional overturning. Considerable variation exists among the OCMIP-2 simulations, with some of the solutions falling noticeably outside available observational constraints. For some cases, model-model and model-data differences can be related to variations in surface forcing, subgrid-scale parameterizations, and model architecture. These errors in the physical metrics point to significant problems in the underlying model representations of ocean transport and dynamics, problems that directly affect the OCMIP predicted ocean tracer and carbon cycle variables (e.g., air-sea CO2 flux, chlorofluorocarbon and anthropogenic CO2 uptake, and export production). A substantial fraction of the large model-model ranges in OCMIP-2 biogeochemical fields (±25–40%) represents the propagation of known errors in model physics. Therefore the model-model spread likely overstates the uncertainty in our current understanding of the ocean carbon system, particularly for transport-dominated fields such as the historical uptake of anthropogenic CO2. A full error assessment, however, would need to account for additional sources of uncertainty such as more complex biological-chemical-physical interactions, biases arising from poorly resolved or neglected physical processes, and climate change.

The Ekman transfer of nutrients and maintenance of new production over the North Atlantic

The maintenance of new production requires a supply of nutrients to the euphotic zone to o⁄set the loss through biological export. The dynamical supply of nutrients is usually discussed in terms of the vertical transfer from nutrient-rich, deep waters. However, the horizontal transfer is important in regions of downwelling over subtropical gyres, where nutrients may be transported across the intergyre boundaries by the surface Ekman drift or geostrophic eddies from the neighbouring nutrient-rich, upwelling regions. The Ekman transfer of nitrate to the euphotic layer is diagnosed from climatology over the North Atlantic. The vertical Ekman supply of nitrate is found to be significant over the subpolar gyre, the tropics and eastern boundary, whereas the horizontal transfer is found to be dominant at the intergyre boundaries. On the northern flank of the subtropical gyre, the Ekman transfer provides a source of nitrate from 0.03 to 0.06 mol N m~2 yr~1, corresponding to a contribution to new production of between 0.4 and 0.8 mol C m~2 yr~1. This estimate represents a significant fraction of the total new production of typically 1 mol C m~2 yr~1 suggested by both remote chlorophyll and sediment trap observations. A simplified nitrogen cycle model is used to assess the role of the Ekman supply over the North Atlantic. In the model the Ekman supply of nitrate leads to a plume of nitrate and enhanced productivity extending up to 1000 km into the subtropical gyre from the intergyre boundaries. This lateral scale is controlled by the seasonal cycle of the mixed layer and the remineralisation of the particulate organic fallout. ( 1998 Elsevier Science Ltd. All rights reserved.

Patterns of Diversity in Marine Phytoplankton

Diversity Gradients Latitudinal gradients in species abundance, with relatively few occurring at the poles and many at the equator, are well known for macroorganisms. It is a matter of controversy, fueled by a lack of observational data, whether such gradients also occur among microorganisms. Barton et al. (p. 1509, published online 25 February) have built on a global marine circulation model to predict the dynamics of phytoplankton populations. In silico, they obtain patterns of latitudinal gradation for plankton that are interspersed with hotspots of amplified diversity, which point to plausible natural explanations for the phenomenon that can be tested in the future by systematic metagenomic surveys. Highest diversity occurs in physically dynamic mid-latitude zones, and lowest diversity and highest biomass occur toward the poles. Spatial diversity gradients are a pervasive feature of life on Earth. We examined a global ocean circulation, biogeochemistry, and ecosystem model that indicated a decrease in phytoplankton diversity with increasing latitude, consistent with observations of many marine and terrestrial taxa. In the modeled subpolar oceans, seasonal variability of the environment led to competitive exclusion of phytoplankton with slower growth rates and lower diversity. The relatively weak seasonality of the stable subtropical and tropical oceans in the global model enabled long exclusion time scales and prolonged coexistence of multiple phytoplankton with comparable fitness. Superimposed on the decline in diversity seen from equator to pole were “hot spots” of enhanced diversity in some regions of energetic ocean circulation, which reflected lateral dispersal.

Decoupling of iron and phosphate in the global ocean

[1] We formulate a mechanistic model of the coupled oceanic iron and phosphorus cycles. The iron parameterization includes scavenging onto sinking particles, complexation with an organic ligand, and a prescribed aeolian source. Export production is limited by the availability of light, phosphate, and iron. We implement this biogeochemical scheme in a coarse resolution ocean general circulation model using scavenging rates and conditional stability constants guided by laboratory studies and a suite of box model sensitivity studies. The model is able to reproduce the broad regional patterns of iron and phosphorus. In particular, the high macronutrient concentrations of the Southern Ocean, tropical Pacific, and subarctic Pacific emerge from the explicit iron limitation of the model. In addition, the model also qualitatively reproduces the observed interbasin gradients of deep, dissolved iron with the lowest values in the Southern Ocean. The ubiquitous presence of significant amounts of free ligand is also explicitly captured. We define a tracer, Fe* which quantifies the degree to which a water mass is iron limited, relative to phosphorus. Surface waters in high-nutrient, lowchlorophyll regions have negative Fe* values, indicating Fe limitation. The extent of the decoupling of iron and phosphorus is determined by the availability and binding strength of the ligand relative to the scavenging by particulate. Global iron concentrations are sensitive to changes in scavenging rate and physical forcing. Decreasing the scavenging rate 40% results in � 0.1 nM increase in dissolved iron in deep waters. Forcing the model with weaker wind stresses leads to a decrease in surface [PO4] and [Fe ]i n the Southern Ocean due to a reduction in the upwelling strength.