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Karl
et al. (1997):
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« Biological
dinitrogen (N2) fixation – the exclusively
prokaryotic metabolic process responsible for converting the most
abundant but relatively inert form of N into biologically
available substrates – is the dominant mechanism for
introduction of N into the biosphere. It approximates the total
losses from microbiological denitrification on the global scale.
Consequently,
data on the quantitative role of N2 fixation in the
oceans’ nutrient budgets are of considerable scientific interest ».
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Recently,
nitrogen fixation and its biogeochemical consequences have
been extensively studied (Gruber and Sarmiento,
1997) and
quantified in the Arabian Sea (Capone et al.,
1998), the open
Atlantic (Carpenter et al., 1999) and Pacific oceans
(Letelier and
Karl, 1996; Karl et al., 1998) and coastal waters
(Bell et al., 1999) N2 fixation is now considered as a significant
input in the marine and global nitrogen cycle (Capone et al.,
1997), and has been introduced into biogeochemical models (Bisset
et al., 1999; Tyrrell, 1999; Walsh et al.,
1999). Tyrrell (1999)
elaborated a simple one-dimentional, two-box model of the global
ocean, which shows how N2 fixation, although low
relative to nitrate input on a global basis, would control on a
long term the global amount of available reactive nitrogen and
therefore the whole ocean productivity.
The
major N2 fixing organisms in Tropical Ocean
In
marine phytoplankton, only some species affiliated to the
phylogenic group of cyanobacteria have the capacity to use
dinitrogen to satisfy their N metabolic requirements. Most
cyanobacteria in tropical and sub-tropical waters belong to 4 main
genus, forming the most fascinating group of marine phytoplankton.
(1) Prochlorococcus are
the smallest and the most numerous oxygenic phototrophic bacteria
(Chisholm et al., 1992) (2) Synechococcus
are fewer and a little larger (Johnson and Sieburth,
1979;
Waterbury et al., 1979) (3) Richelia
lives in symbiosis in some species of diatoms such as Rhizosolenia
(Burford et al., 1995) or Hemiaulus
(Villareal, 1991) Richelia
form filamentous colonies called trichomes and have heterocysts
which are special cells fixing nitrogen. (4) Trichodesmium
also form trichomes but have no heterocyst. Trichomes are often
associated as spherical aggregates called puffs or as bundles
called tufts. Trichodesmium, though lacking heterocysts, are known to fix nitrogen
(Dugdale et al., 1961) They are probably the most quantitatively
important cyanobacteria in terms of oceanic N input. By contrast,
experiments intended to demonstrate N2 fixation in
natural Prochlorococcus and Synechococcus
are not yet conclusive to date, even if genetic analyses do not
exclude such a possibility (Zehr et al.,
1998).
N2
fixation in the oceans: magnitude underestimated
Quantitative
studies devoted to N2 fixation have proved that at the
basin scale, the rate of N2 fixation in the nitrogen
balance has been largely underestimated (Gruber and Sarmiento,
1997), in part owing to spatial and temporal undersampling of the
marine environment. Most historical estimates have shown low rates
of N2 fixation relative to the total N requirement for
primary production in the sea (Karl et al.,
1997) By contrast,
several recent studies demonstrate the contrary. For example, Karl
et al. (1997) have reported that at station ALOHA north of Hawaii,
N2 fixation could satisfy up to 50% of nitrogen
requirements needed to sustain the export of particulate matter
out of the upper layer. In other words, input of N by diazotrophic
activity would represent an important source of « new »
nitrogen (sensu Dugdale
and Goering, 1967) at ALOHA station but probably also in the whole
north and south central gyres, especially during El Niño periods
(Karl et al., 1997) .
Furthermore,
recent geochemical studies indicate that N2 fixation
rates in the North Atlantic are at least twice as high as rates
derived from direct measurements (Hood et al.,
1999).
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