David M. Glover

Iron cycling and nutrient-limitation patterns in surface waters of the World Ocean

Abstract A global marine ecosystem mixed-layer model is used to study iron cycling and nutrient-limitation patterns in surface waters of the world ocean. The ecosystem model has a small phytoplankton size class whose growth can be limited by N, P, Fe, and/or light, a diatom class which can also be Si-limited, and a diazotroph phytoplankton class whose growth rates can be limited by P, Fe, and/or light levels. The model also includes a parameterization of calcification by phytoplankton and is described in detail by Moore et al. (Deep-Sea Res. II, 2002). The model reproduces the observed high nitrate, low chlorophyll (HNLC) conditions in the Southern Ocean, subarctic Northeast Pacific, and equatorial Pacific, and realistic global patterns of primary production, biogenic silica production, nitrogen fixation, particulate organic carbon export, calcium carbonate export, and surface chlorophyll concentrations. Phytoplankton cellular Fe/C ratios and surface layer dissolved iron concentrations are also in general agreement with the limited field data. Primary production, community structure, and the sinking carbon flux are quite sensitive to large variations in the atmospheric iron source, particularly in the HNLC regions, supporting the Iron Hypothesis of Martin (Paleoceanography 5 (1990) 1–13). Nitrogen fixation is also strongly influenced by atmospheric iron deposition. Nitrogen limits phytoplankton growth rates over less than half of the world ocean during summer months. Export of biogenic carbon is dominated by the sinking particulate flux, but detrainment and turbulent mixing account for 30% of global carbon export. Our results, in conjunction with other recent studies, suggest the familiar paradigm that nitrate inputs to the surface layer can be equated with particulate carbon export needs to be expanded to include multiple limiting nutrients and modes of export.

An intermediate complexity marine ecosystem model for the global domain

A new marine ecosystem model designed for the global domain is presented, and model output is compared with field data from nine different locations. Field data were collected as part of the international Joint Global Ocean Flux Study (JGOFS) program, and from historical time series stations. The field data include a wide variety of marine ecosystem types, including nitrogen- and iron-limited systems, and different physical environments from high latitudes to the mid-ocean gyres. Model output is generally in good agreement with field data from these diverse ecosystems. These results imply that the ecosystem model presented here can be reliably applied over the global domain. The model includes multiple potentially limiting nutrients that regulate phytoplankton growth rates. There are three phytoplankton classes, diatoms, diazotrophs, and a generic small phytoplankton class. Growth rates can be limited by available nitrogen, phosphorus, iron, and/or light levels. The diatoms can also be limited by silicon. The diazotrophs are capable of nitrogen fixation of N2 gas and cannot be nitrogen-limited. Calcification by phytoplankton is parameterized as a variable fraction of primary production by the small phytoplankton group. There is one zooplankton class that grazes the three phytoplankton groups and a large detrital pool. The large detrital pool sinks out of the mixed layer, while a smaller detrital pool, representing dissolved organic matter and very small particulates, does not sink. Remineralization of the detrital pools is parameterized with a temperature-dependent function. We explicitly model the dissolved iron cycle in marine surface waters including inputs of iron from subsurface sources and from atmospheric dust deposition.

A new coupled, one-dimensional biological-physical model for the upper ocean: Applications to the JGOFS Bermuda Atlantic Time-series Study (BATS) site

Abstract This paper presents a new coupled, one dimensional biological-physical model applied to the subtropical region near Bermuda. The physical component of the model, which is driven by smooth climatological forcing, successfully reproduces the long-term seasonal cycles of upper ocean temperature, salinity and boundary layer depth from Hydrostation S. The nitrogen-based biological model, which includes the effects of photoadaptation, phytoplankton aggregation, and particle remineralization in the aphotic zone, shows significant skill in capturing the major features of the annual chlorophyll field (e.g. spring bloom, deep chlorophyll maximum) and depth-integrated chlorophyll and primary production as exhibited by the U.S. JGOFS Bermuda Atlantic Time-series Study (BATS) data. The introduction of variable phytoplankton chlorophyll-to-nitrogen ratios is found to be important for simulating the subsurface chlorophyll maximum, and the model solutions show a realistic deep nitracline in the summer and a low annual average f -ratio of ∼0.21 compared to previous modeling work. The performance of the model solutions are weakest during the late summer, when the model can not supply enough nutrients to support the high production observed in the stratified near-surface waters. The coupled model has large winter production values, leading to a substantial export of organic material from the euphotic zone via downward turbulent mixing. The model predicts a total export production from the euphotic zone of 0.24 mol N m −2 year −1 , approximately equally partitioned between particle sinking and suspended matter detrainment. The bulk of the export production is remineralized in the shallow aphotic zone, and only a small fraction is transported below the depth of the maximum winter mixed layer and thus contributes to “biological pump”.

Estimates of wintertime mixed layer nutrient concentrations in the North Atlantic

Abstract Nonlinear, time-dependent model sensitivity to initial conditions poses a challenging problem when attempting to initialize such a model. In order to intialize a chemical-physical model of the upper several hundred meters of the North Atlantic, we have calculated the initial concentrations of several chemical species from three estimation methods by a combination of the Climatological Atlas of the World Ocean ( Levitus , 1982) and the TTO north, and tropical, Atlantic study data bases. A 1° × 1° grid of the average initial concentrations over the mixed layer depth was generated for the method of preference and added to the inialization data base of the model. Contour maps of this calculated initial concentration set are presented and comparisons with the other methods and actual data are made.

A model function of the global bomb tritium distribution in precipitation, 1960–1986

We have developed, using the World Meteorological Organization/International Atomic Energy Agency (WMO/IAEA) tritium data set, a global model function for predicting the annual mean concentration of decay-corrected bomb tritium (TU81N) in precipitation over the time period 1960–1986. The model consists of two reference time histories or factors, calculated from a factor analysis of the zonally averaged global data set, and global maps of the two spatial coefficents associated with the factors. By combining the reference curves with the appropriate coefficent values taken from these maps, an estimate of the tritium time history for particular locations can be produced. The predicted error for the integrated tritium concentration from the annual model is approximately 3–10% and, in a comparison with previous work (Weiss and Roether, 1980), provides a statistically better fit of the tritium data at most stations. We also discuss the seasonal cycle of tritium in precipitation and present estimates for the amplitude and phase for tritium using a simple seasonal model. At most of the WMO/IAEA monitoring stations, our four-parameter seasonal model accounts for greater than 80% of the variance in the monthly observations.

The pH of the North Atlantic Ocean: Improvements to the global model for sound absorption in seawater

At frequencies below 1 kHz, sound absorption coefficients in the ocean are a function of pH, and at higher frequencies they are dependent upon MgSO4. The pH dependent terms are attributable to relaxation of B(OH)3 and MgCO3 species, and the ensemble effect has been approximated (Mellen et al., 1987a) as α = α1(MgSO4) + α2(B(OH)3) + α3(MgCO3), where α is the absorption coefficient in decibels per kilometer, and αn = (S/35)An[ƒ2ƒn/(ƒ2 + ƒn2)], where S is the salinity, An is the amplitude of each component and depends on pH, ƒ is the frequency, and ƒn is the relaxation frequency. Overall accuracy of ±15% in a requires that pH be known to 0.05 pH units. The presently used oceanic pH field for sound absorption models is derived from a combination of Geochemical Ocean Sections Study (GEOSECS) data and Soviet data from the 1978 Gorshkov atlas for the North Atlantic where GEOSECS data are absent. We compare the North Atlantic fields with the well-constrained Transient Tracers in the Ocean North Atlantic CO2 data set and find large differences. We further show that sufficiently strong correlations exist between CO2 system variables and other more commonly available hydrographic properties and that improved renditions of the pH field in other regions of the ocean are possible once equivalent local correlations are established, thus leading to greatly improved estimates of the sound absorption field.

Dynamics of the transition zone in coastal zone color scanner-sensed ocean color in the North Pacific during oceanographic spring

A transition zone in phytoplankton concentration running across the North Pacific basin at 30° to 40° north latitude corresponds to a basin-wide front in surface chlorophyll observed in a composite of coastal zone color scanner (CZCS) images for May, June, and July 1979–1986. This transition zone with low chlorophyll to the south and higher chlorophyll to the north can be simulated by a simple model of the concentration of phytoplankton, zooplankton, and dissolved nutrient (nitrate) in the surface mixed layer of the ocean applied to the North Pacific basin for the climatological conditions during oceanographic springtime (May, June, and July). The model is initialized with a 1° × 1° gridded estimate of wintertime (February, March, and April) mixed layer nitrate concentrations calculated from an extensive nutrient database and a similarly gridded mixed layer depth data set. Comparison of model predictions with CZCS data provides a means of evaluating the dynamics of the transition zone. We conclude that in the North Pacific, away from major boundary currents and coastal upwelling zones, wintertime vertical mixing determines the total nutrient available to the plankton ecosystem in the spring. The transition zone seen in basin-scale CZCS images is a reflection of the geographic variation in the wintertime mixed layer depth and the nitracline, leading to a latitudinal gradient in phytoplankton chlorophyll.

Life-cycle modification in open oceans accounts for genome variability in a cosmopolitan phytoplankton.

Emiliania huxleyi is the most abundant calcifying plankton in modern oceans with substantial intraspecific genome variability and a biphasic life cycle involving sexual alternation between calcified 2N and flagellated 1N cells. We show that high genome content variability in Emiliania relates to erosion of 1N-specific genes and loss of the ability to form flagellated cells. Analysis of 185 E. huxleyi strains isolated from world oceans suggests that loss of flagella occurred independently in lineages inhabiting oligotrophic open oceans over short evolutionary timescales. This environmentally linked physiogenomic change suggests life cycling is not advantageous in very large/diluted populations experiencing low biotic pressure and low ecological variability. Gene loss did not appear to reflect pressure for genome streamlining in oligotrophic oceans as previously observed in picoplankton. Life-cycle modifications might be common in plankton and cause major functional variability to be hidden from traditional taxonomic or molecular markers.

An Integrated Assessment Model for Helping the United States Sea Scallop (Placopecten magellanicus) Fishery Plan Ahead for Ocean Acidification and Warming

Ocean acidification, the progressive change in ocean chemistry caused by uptake of atmospheric CO2, is likely to affect some marine resources negatively, including shellfish. The Atlantic sea scallop (Placopecten magellanicus) supports one of the most economically important single-species commercial fisheries in the United States. Careful management appears to be the most powerful short-term factor affecting scallop populations, but in the coming decades scallops will be increasingly influenced by global environmental changes such as ocean warming and ocean acidification. In this paper, we describe an integrated assessment model (IAM) that numerically simulates oceanographic, population dynamic, and socioeconomic relationships for the U.S. commercial sea scallop fishery. Our primary goal is to enrich resource management deliberations by offering both short- and long-term insight into the system and generating detailed policy-relevant information about the relative effects of ocean acidification, temperature rise, fishing pressure, and socioeconomic factors on the fishery using a simplified model system. Starting with relationships and data used now for sea scallop fishery management, the model adds socioeconomic decision making based on static economic theory and includes ocean biogeochemical change resulting from CO2 emissions. The model skillfully reproduces scallop population dynamics, market dynamics, and seawater carbonate chemistry since 2000. It indicates sea scallop harvests could decline substantially by 2050 under RCP 8.5 CO2 emissions and current harvest rules, assuming that ocean acidification affects P. magellanicus by decreasing recruitment and slowing growth, and that ocean warming increases growth. Future work will explore different economic and management scenarios and test how potential impacts of ocean acidification on other scallop biological parameters may influence the social-ecological system. Future empirical work on the effect of ocean acidification on sea scallops is also needed.

Using altimetry to help explain patchy changes in hydrographic carbon measurements

Here we use observations and ocean models to identify mechanisms driving large seasonal to interannual variations in dissolved inorganic carbon (DIC) and dissolved oxygen (O-2) in the upper ocean. We begin with observations linking variations in upper ocean DIC and O-2 inventories with changes in the physical state of the ocean. Models are subsequently used to address the extent to which the relationships derived from short-timescale (6 months to 2 years) repeat measurements are representative of variations over larger spatial and temporal scales. The main new result is that convergence and divergence (column stretching) attributed to baroclinic Rossby waves can make a first-order contribution to DIC and O-2 variability in the upper ocean. This results in a close correspondence between natural variations in DIC and O-2 column inventory variations and sea surface height (SSII) variations over much of the ocean. Oceanic Rossby wave activity is an intrinsic part of the natural variability in the climate system and is elevated even in the absence of significant interannual variability in climate mode indices. The close correspondence between SSII and both DIC and O-2 column inventories for many regions suggests that SSII changes (inferred from satellite altimetry) may prove useful in reducing uncertainty in separating natural and anthropogenic DIC signals (using measurements from Climate Variability and Predictabilitys CO2/Repeat Hydrography program).