A basic problem in marine biogeochemistry is understanding material and elemental distributions and fluxes in the oceans, and a key part of this problem is understanding the processes that affect particulate material in the ocean. Aggregation of particulate material is a primary process because it alters the transport properties of particulate material and provides a mechanism for transferring material from the dissolved into the particulate pools. Aggregation theory not only provides a framework for understanding these processes, but it also provides a means for making predictions and has been successfully used to predict maximum particle concentrations in the oceans and the fate of diatom blooms (including those from iron fertilization), the size spectra of particles in the oceans, and the size distributions of trace metals. Here we review the basic theory involved, summarize recent developments, and explore unresolved issues.

Autotrophic picoplankton dominate primary production over large oceanic regions but are believed to contribute relatively little to carbon export from surface layers. Using analyses of data from the equatorial Pacific Ocean and Arabian Sea, we show that the relative direct and indirect contribution of picoplankton to export is proportional to their total net primary production, despite their small size. We suggest that all primary producers, not just the large cells, can contribute to export from the surface layer of the ocean at rates proportional to their production rates.

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We recorded vertical profiles of size distributions of large particles (.100 mm) to a 1000-m depth in the Atlantic, Indian, and Pacific Oceans and in the Mediterranean Sea with the Underwater Video Profiler. Of the 410 profiles used in our analysis, 193 also included temperature, salinity, and high-performance liquid chromatography (HPLC)-resolved pigments, which were used to characterize the size structure of the phytoplankton community. Classification analysis identified six clusters of vertical profiles of size distributions of particles. Each cluster was characterized by the size distribution of its particles in the mesopelagic layer and the change of the particle-size distribution with depth. Clusters with large particles in the mesopelagic layer corresponded to surface waters dominated by microphytoplankton, and those with small particles corresponded to surface waters dominated by picophytoplankton. We estimated the mass flux at 400 m using a relationship between particle size and mass flux. Principal-component regression analysis showed that 68% of the variance of the mass flux at 400 m was explained by the size structure of the phytoplankton community and integrated chlorophyll a in the euphotic zone. We found that coefficient k in the Martin power relationship, which describes the decrease in the vertical mass flux with depth, varies between 0.2 and 1.0 in the world ocean, and we provided an empirical relationship to derive k from the size structure of phytoplankton biomass in the euphotic zone. Biogeochemists and modelers could use that relationship to obtain a realistic description of the downward particle flux instead of using a constant k value as often done.

Leaking organic solutes form an elongated plume in the wake of a sinking aggregate. These solutes may both be assimilated by suspended bacteria and guide bacteria with chemokinetic swimming behavior toward the aggregate. We used modifications of previously published models of the flow and concentration fields around sinking aggregates and of chemokinetic behavior of bacteria to identify the behavior that optimizes aggregate colonization and plume utilization. The optimal solution is governed by physical constraints and is a trade off between a high sensitivity to chemical signals and a long signal integration time. For a run-and-tumble swimming behavior, the predicted tumbling frequency is between 1 and 10 s Ϫ1 , similar to that reported for marine bacteria. The predicted optimal sensitivity to chemical signals is similar to or greater than that known for Escherichia coli. The optimal behavior was used to examine the potential contribution of aggregate-generated solute plumes for water column bacterial production. Despite occupying only a small volume fraction, the plumes may provide important growth habitats for free bacteria and account for a significant proportion of water column bacterial production at typical concentrations of marine snow aggregates.

a b s t r a c tMetabolic activity in the water column below the euphotic zone is ultimately fuelled by the vertical flux of organic material from the surface. Over time, the deep ocean is presumably at steady state, with sources and sinks balanced. But recently compiled global budgets and intensive local field studies suggest that estimates of metabolic activity in the dark ocean exceed the influx of organic substrates. This imbalance indicates either the existence of unaccounted sources of organic carbon or that metabolic activity in the dark ocean is being over-estimated. Budgets of organic carbon flux and metabolic activity in the dark ocean have uncertainties associated with environmental variability, measurement capabilities, conversion parameters, and processes that are not well sampled. We present these issues and quantify associated uncertainties where possible, using a Monte Carlo analysis of a published data set to determine the probability that the imbalance can be explained purely by uncertainties in measurements and conversion factors. A sensitivity analysis demonstrates that the bacterial growth efficiencies and assumed cell carbon contents have the greatest effects on the magnitude of the carbon imbalance. Two poorly quantified sources, lateral advection of particles and a population of slowly settling particles, are discussed as providing a means of closing regional carbon budgets. Finally, we make recommendations concerning future research directions to reduce important uncertainties and allow a better determination of the magnitude and causes of the unbalanced carbon budgets.

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