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leading to overexploitation. Fishing practices such as the use of drift nets in the North Pacific
Ocean may also have devastating effects on marine stocks and ecosystems. In addition, the
increasing frequency of exceptional phytoplankton blooms (e.g. red tides, brown tides, green
tides, etc.) may imperil coastal fisheries and aquaculture operations (e.g. White, 1984;
Gosselin et al., 1989). Recent reviews (e.g. Parsons and Lalli, 1988; Cushing, 1989;
Legendre, 1990) have suggested that one of the key factors in the control of exploited marine
populations may be phytoplankton production and its export pathways.
Flow cytometry and image analysis, which provide estimates of particle size together with
simultaneous cell-by-cell measurements of various characteristics, are playing an increasingly
important role in biological oceanography (see the review by Legendre and Yentsch, 1989).
So far, a large share of the effort has been devoted to the development of techniques suitable
for analyzing aquatic particles. As suggested by Legendre and Yentsch (1989), the next phase,
which is already under way in several laboratories, will be to use these techniques for
developing and testing theories in biological oceanography. Within this context, the present
paper will show that a significant component of the biologically driven flux of carbon dioxide
into the oceans is determined from the level of individual cells up, and mediated by ecosystem
properties. This is significant for the sequestration of carbon in the oceans and also for
exploited renewable marine resources. It will be concluded that relevant characteristics of
biogenic particle dynamics and of marine ecosystems must be incorporated into the models
developed for the management of exploited stocks and for the prediction of the global flux of
carbon into the oceans 4 •
ROLE OF PELAGIC MARINE ECOSYSTEMS IN THE SEQUESTRATION OF
CARBON
It is often assumed, at least implicitly, that the biological CO 2 pump is essentially driven by
the upward flux of nitrate, so that the potential export of biogenic carbon from the upper
ocean would be stoichiometrically equivalent to phytoplankton new production (e.g. Eppley
and Peterson, 1979), as estimated from the uptake of 15N03- (Dugdale and Goering, 1967) or
the upward flux of N03- (e.g. Lewis et al., 1986). This approach assumes that ecosystems are
4 Table I gives glossary of some oceanographic terms, as understood in the present paper.
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