Transparent exopolymer particles (TEP) in aquatic environments.

A pelagic food web model was formulated with the goal of developing a quantitative understanding of the relationship between total production, export production, and environmental variables in marine ecosystems. The model assumes that primary production is partitioned through both large and small phytoplankton and that the food web adjusts to changes in the rate of allochthonous nutrient inputs in a way that maximizes stability, i.e., the ability of the system to return to steady state following a perturbation. The results of the modeling exercise indicate that ef ratios, defined as new production/total production = export production/total production, are relatively insensitive to total production rates at temperatures greater than ∼25°C and lie in the range 0.1-0.2. At moderate to high total production rates, ef ratios are insensitive to total production and negatively correlated with temperature. The maximum ef ratios are ∼0.67 at high rates of production and temperatures of 0°−10°C. At temperatures less than ∼20°C, there is a transition from low ef ratios to relatively high ef ratios as total production increases from low to moderate values. This transition accounts for the hyperbolic relationship often presumed to exist between ef ratios and total production. At low rates of production the model predicts a negative correlation between production and ef ratios, a result consistent with data collected at station ALOHA (22°45′N, 158°W) in the North Pacific subtropical gyre. The predictions of the model are in excellent agreement with results reported from the Joint Global Ocean Flux Study (JGOFS) and from other field work. In these studies, there is virtually no correlation between total production and ef ratios, but temperature alone accounts for 86% of the variance in the ef ratios. Model predictions of the absolute and relative abundance of autotrophic and heterotrophic microorganisms are in excellent agreement with data reported from field studies. Combining the ef ratio model with estimates of ocean temperature and photosynthetic rates derived from satellite data indicates that export production on a global scale is ∼20% of net photosynthesis. The results of the model have important implications for the impact of climate change on export production, particularly with respect to temperature effects.

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Temperature effects on export production in the open ocean

[1] Mid-Cretaceous (Barremian-Turonian) plankton preserved in deep-sea marl, organic-rich shale, and pelagic carbonate hold an important record of how the marine biosphere responded to short- and long-term changes in the ocean-climate system. Oceanic anoxic events (OAEs) were short-lived episodes of organic carbon burial that are distinguished by their widespread distribution as discrete beds of black shale and/or pronounced carbon isotopic excursions. OAE1a in the early Aptian (∼120.5 Ma) and OAE2 at the Cenomanian/Turonian boundary (∼93.5 Ma) were global in their distribution and associated with heightened marine productivity. OAE1b spans the Aptian/Albian boundary (∼113–109 Ma) and represents a protracted interval of dysoxia with multiple discrete black shales across parts of Tethys (including Mexico), while OAE1d developed across eastern and western Tethys and in other locales during the latest Albian (∼99.5 Ma). Mineralized plankton experienced accelerated rates of speciation and extinction at or near the major Cretaceous OAEs, and strontium isotopic evidence suggests a possible link to times of rapid oceanic plateau formation and/or increased rates of ridge crest volcanism. Elevated levels of trace metals in OAE1a and OAE2 strata suggest that marine productivity may have been facilitated by increased availability of dissolved iron. The association of plankton turnover and carbon isotopic excursions with each of the major OAEs, despite the variable geographic distribution of black shale accumulation, points to widespread changes in the ocean-climate system. Ocean crust production and hydrothermal activity increased in the late Aptian. Faster spreading rates [and/or increased ridge length] drove a long-term (Albian–early Turonian) rise in sea level and CO2-induced global warming. Changes in ocean circulation, water column stratification, and nutrient partitioning lead to a reorganization of plankton community structure and widespread carbonate (chalk) deposition during the Late Cretaceous. We conclude that there were important linkages between submarine volcanism, plankton evolution, and the cycling of carbon through the marine biosphere.

Oceanic anoxic events and plankton evolution: Biotic response to tectonic forcing during the mid-Cretaceous

Ecosystem processes are important determinants of the biogeochemistry of the ocean, and they can be profoundly affected by changes in climate. Ocean models currently express ecosystem processes through empirically derived parameterizations that tightly link key geochemical tracers to ocean physics. The explicit inclusion of ecosystem processes in models will permit ecological changes to be taken into account, and will allow us to address several important questions, including the causes of observed glacial-interglacial changes in atmospheric trace gases and aerosols, and how the oceanic uptake of CO2 is likely to change in the future. There is an urgent need to assess our mechanistic understanding of the environmental factors that exert control over marine ecosystems, and to represent their natural complexity based on theoretical understanding. We present a prototype design for a Dynamic Green Ocean Model (DGOM) based on the identification of (a) key plankton functional types that need to be simulated explicitly to capture important biogeochemical processes in the ocean; (b) key processes controlling the growth and mortality of these functional types and hence their interactions; and (c) sources of information necessary to parameterize each of these processes within a modeling framework. We also develop a strategy for model evaluation, based on simulation of both past and present mean state and variability, and identify potential sources of validation data for each. Finally, we present a DGOM-based strategy for addressing key questions in ocean biogeochemistry. This paper thus presents ongoing work in ocean biogeochemical modeling, which, it is hoped will motivate international collaborations to improve our understanding of the role of the ocean in the climate system.

Ecosystem dynamics based on plankton functional types for global ocean biogeochemistry models

Predation in aquatic microbial food webs is dominated by phagotrophic protists, yet these microorganisms are still understudied compared to bacteria and phytoplankton. In pelagic ecosystems, predaceous protists are ubiquitous, range in size from 2 µm flagellates to >100 µm ciliates and dinoflagellates, and exhibit a wide array of feeding strategies. Their trophic states run the gamut from strictly phagotrophic, to mixotrophic: partly autotrophic and partly phagotrophic, to primarily autotrophic but capable of phagotrophy. Protists are a major source of mortality for both heterotrophic and autotrophic bacteria. They compete with herbivorous meso- and macro-zooplankton for all size classes of phytoplankton. Protist grazing may affect the rate of organic sinking flux from the euphotic zone. Protist excretions are an important source of remineralized nutrients, and of colloidal and dissolved trace metals such as iron, in aquatic systems. Work on predation by protists is being facilitated by methodological advances, e.g., molecular genetic analysis of protistan diversity and application of flow cytometry to study population growth and feeding rates. Examples of new research areas are studies of impact of protistan predation on the community structure of prey assemblages and of chemical communication between predator and prey in microbial food webs.

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Significance of predation by protists in aquatic microbial food webs.

Models of biogenic carbon (BC) flux assume that short herbivorous food chains lead to high export, whereas complex microbial or omnivorous food webs lead to recycling and low export, and that export of BC from the euphotic zone equals new production (NP). In the Gulf of St. Lawrence, particulate organic carbon fluxes were similar during the spring phytoplankton bloom, when herbivory dominated, and during nonbloom conditions, when microbial and omnivorous food webs dominated. In contrast, NP was 1.2 to 161 times greater during the bloom than after it. Thus, neither food web structure nor NP can predict the magnitude or patterns of BC export, particularly on time scales over which the ocean is in nonequilibrium conditions.

Kenneth Crocker

Papers

Vertical Flux of Biogenic Carbon in the Ocean: Is There Food Web Control?