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	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>5</volume_number>
		<issue_number>2</issue_number>
		<publication_year>2008</publication_year>
	</journal>
	<doi>10.5194/bg-5-323-2008</doi>
	<article_url>http://www.biogeosciences.net/5/323/2008/</article_url>
	<abstract_html>http://www.biogeosciences.net/5/323/2008/bg-5-323-2008.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/5/323/2008/bg-5-323-2008.pdf</fulltext_pdf>
	<start_page>323</start_page>
	<end_page>338</end_page>
	<publication_date>2008-03-05</publication_date>
	<article_title content_type="html">Evidence for efficient regenerated production and dinitrogen fixation in nitrogen-deficient waters of the South Pacific Ocean: impact on new and export production estimates</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>P. Raimbault</name>
			<email>patrick.raimbault@univmed.fr</email>
		</author>
		<author numeration="2" affiliations="1">
			<name>N. Garcia</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Laboratoire de Microbiologie, Géochimie et Ecologie Marine (UMR 6117 CNRS), Centre d&apos;Océanologie de Marseille, Université de la Méditerranée, Campus de Luminy, 13288 Marseille, France</affiliation>
	</affiliations>
	<abstract content_type="html">One of the major objectives of the BIOSOPE cruise, carried out on the R/V
Atalante from October-November 2004 in the South Pacific Ocean, was to
establish productivity rates along a zonal section traversing the
oligotrophic South Pacific Gyre (SPG). These results were then compared to
measurements obtained from the nutrient &amp;ndash; replete waters in the Chilean
upwelling and around the Marquesas Islands. A dual &lt;sup&gt;13&lt;/sup&gt;C/&lt;sup&gt;15&lt;/sup&gt;N isotope
technique was used to estimate the carbon fixation rates, inorganic nitrogen
uptake (including dinitrogen fixation), ammonium (NH&lt;sub&gt;4&lt;/sub&gt;) and nitrate
(NO&lt;sub&gt;3&lt;/sub&gt;) regeneration and  release of dissolved organic nitrogen (DON). The SPG
exhibited the lowest primary production rates (0.15 g C m&lt;sup&gt;&amp;minus;2&lt;/sup&gt; d&lt;sup&gt;&amp;minus;1&lt;/sup&gt;),
while rates were 7 to 20 times higher around the Marquesas Islands and in
the Chilean upwelling, respectively. In the very low productive area of the
SPG, most of the primary production was sustained by active regeneration
processes that fuelled up to 95% of the biological nitrogen demand.
Nitrification was active in the surface layer and often balanced the
biological demand for nitrate, especially in the SPG. The percentage of
nitrogen released as DON represented a large proportion of the inorganic
nitrogen uptake (13&amp;ndash;15% in average), reaching 26&amp;ndash;41% in the SPG, where
DON production played a major role in nitrogen cycling. Dinitrogen fixation
was detectable over the whole study area; even in the Chilean upwelling, where
rates as high as 3 nmoles l&lt;sup&gt;&amp;minus;1&lt;/sup&gt; d&lt;sup&gt;&amp;minus;1&lt;/sup&gt; were measured. In these
nutrient-replete waters new production was very high (0.69&amp;plusmn;0.49 g C m&lt;sup&gt;&amp;minus;2&lt;/sup&gt; d&lt;sup&gt;&amp;minus;1&lt;/sup&gt;)
 and essentially sustained by nitrate levels. In the
SPG, dinitrogen fixation, although occurring at much lower daily rates
(&amp;asymp;1&amp;ndash;2 nmoles l&lt;sup&gt;&amp;minus;1&lt;/sup&gt; d&lt;sup&gt;&amp;minus;1&lt;/sup&gt;), sustained up to 100% of the
new production (0.008&amp;plusmn;0.007 g C m&lt;sup&gt;&amp;minus;2&lt;/sup&gt; d&lt;sup&gt;&amp;minus;1&lt;/sup&gt;) which was two
orders of magnitude lower than that measured in the upwelling. The annual
N&lt;sub&gt;2&lt;/sub&gt;-fixation of the South Pacific is estimated to 21&amp;times;10&lt;sup&gt;12&lt;/sup&gt;g,
of which 1.34&amp;times;10&lt;sup&gt;12&lt;/sup&gt;g is for the SPG only. Even if our &quot;snapshot&quot;
estimates of N&lt;sub&gt;2&lt;/sub&gt;-fixation rates were lower than that expected from a
recent ocean circulation model, these data confirm that the N-deficiency
South Pacific Ocean would provide an ideal ecological niche for the
proliferation of N&lt;sub&gt;2&lt;/sub&gt;-fixers which are not yet identified.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Allen, A. E, Howard-Jones, M. H., Booth, M. G., Frischer, M. E., Verity, P. G., Bronk, D. A., Sanderson, M. P: Importance of heterotrophic bacterial assimilation of ammonium and nitrate in the Barents Sea during summer, J. Marine Syst., 38, 93&amp;ndash;108, 2002. </reference>
		<reference numeration="2" content_type="text"> Altabet, M.: Organic C, N and stable isotopic composition of particulate matter collected on glass-fibber and aluminum oxide filters, Limnol. Oceanogr., 35(4), 902&amp;ndash;909, 1990. </reference>
		<reference numeration="3" content_type="text"> Aufdenkampe, A. K., McCarthy, J. J., Rodier, M., Navarrette, C., Dunne, J., and Murray, J. W.: Estimation of new production in the Tropical Pacific, Global Biogeochem. Cy., 15, 101&amp;ndash;113, 2001. </reference>
		<reference numeration="4" content_type="text"> Aufdenkampe, A. K., McCarthy, J. J., Navarrette, C., Rodier, M., Dunne, J., and Murray, J. W.: Biogeochemical controls on new production in the tropical pacific, Deep. Sea. Res. II, 49, 2619&amp;ndash;2648, 2002. </reference>
		<reference numeration="5" content_type="text"> Béthoux, J. P. and Copin-Montégut, G.: Biological fixation of atmospheric nitrogen in the Mediterranean Sea, Limnol. Oceanogr., 31, 1353&amp;ndash;1358, 1986. </reference>
		<reference numeration="6" content_type="text"> Bianchi, M., Feliatra, F., Tréguer, P., Vincendeau, M., and Morvan, J.: Nitrification rates, ammonium and nitrate distribution in upper layers of the water column and in sediments of the Indian sector of the Southern Ocean, Deep Sea Res. II, 44(5), 1017&amp;ndash;1032, 1997. </reference>
		<reference numeration="7" content_type="text"> Biegala, I. C. and Raimbault, P.: High abundance of diazotrophic picocyanobacteria (&amp;lt;3 μm) in a South-West Pacific coral lagoon, Aquat. Microb. Ecol., in press, 2008. </reference>
		<reference numeration="8" content_type="text"> Bode, A., Castro, C., Doval, M. D., and Varela, M.: New and regenerated production and ammonium regeneration in the western Bransfield region (Antarctica) during phytoplankton bloom conditions in summer, Deep Sea Res. II, 49, 787&amp;ndash;804, 2002. </reference>
		<reference numeration="9" content_type="text"> Bronk, D. A. and Glibert, P.: A $^15$N tracer method for the measurement of dissolved organic nitrogen release by phytoplankton, Mar. Ecol. Prog. Ser., 77, 171&amp;ndash;182 1991. </reference>
		<reference numeration="10" content_type="text"> Bronk, D. A., Glibert, P., and Ward, B. B.: Nitrogen uptake, dissolved organic nitrogen release and new production, Science, 256, 1843&amp;ndash;1846, 1994. </reference>
		<reference numeration="11" content_type="text"> Capone, D. G. and Carpenter, E. J.: Nitrogen fixation in the marine environment, Science, 217, 1140&amp;ndash;1142, 1982. </reference>
		<reference numeration="12" content_type="text"> Capone, D. G., Burns, J. A, Montoya, J. P., Subramaniam, A., Mahaffey, C., Gunderson, T., Michaels, A. F., and Carpenter, E. J.: Nitrogen fixation by Trichodesmium spp.: An important source of new nitrogen to the tropical and subtropical North Atlantic Ocean, Global Biogeochem. Cy., 19, GB2024, doi:10.1029/2004GB002331, 2005. </reference>
		<reference numeration="13" content_type="text"> Carlson, C. A., Ducklow, H. W., and Michaels, A. F.: Annual flux of dissolved organic carbon from the euphotic zone in the northwestern Sargasso Sea, Nature, 371, 405&amp;ndash;408, 1994. </reference>
		<reference numeration="14" content_type="text"> Chavez, F. P., Buck, K. R., Bidigare, R. R., Karl, D. M., Hebel, D., Latasa, M., and Campbell, L.: On the chlorophyll a retention properties of glass-fiber GF/F filters, Limnol. Oceanogr., 40, 428&amp;ndash;433, 1995. </reference>
		<reference numeration="15" content_type="text"> Claustre, H. and Maritorena, S.: Ocean Science: The many shades of ocean blue, Science, 302, 1514&amp;ndash;1515, 2003. </reference>
		<reference numeration="16" content_type="text"> Codispoti, L. A. and Christensen, J. P.: Nitrification, denitrification and nitrous oxide cycling in the eastern tropical South Pacific Ocean, Mar. Chem., 16(4), 277&amp;ndash;300, 1985. </reference>
		<reference numeration="17" content_type="text"> Copin-Montégut, G. and Avril, B.: Vertical distribution and temporal varaition of dissolved organic carbon in the North-Western Mediterranean Sea, Deep-Sea Res., 40, 1963&amp;ndash;1972, 1993. </reference>
		<reference numeration="18" content_type="text"> Daneri, G. and Quinones, R. A.: Undersamples ocean systems: appeal for an international study of biogeochemical cycles in the Southern Pacific gyre and its boundaries, U.S. JGOFS Newsletter January 2001, 9, 2001. </reference>
		<reference numeration="19" content_type="text"> Deutsch, C., Sarmiento, J.L, Sigman, D. M., Gruber, N., and Dunne, J. P.: Spatial coupling of nitrogen inputs and losses in the ocean, 445(7124), 163&amp;ndash;167, 2007. </reference>
		<reference numeration="20" content_type="text"> Diaz, F. and Raimbault, P.: Nitrogen regeneration and dissolved organic nitrogen release during spring in a NW Mediterranean coastal zone (Gulf of Lions): implications for the estimations of f-ratio and new production, Mar. Ecol. Progr. Ser., 197, 53&amp;ndash;66, 2000. </reference>
		<reference numeration="21" content_type="text"> Dore, J. E. and Karl, D. M.: Nitrification in the euphotic zone as a source of nitrite, nitrate, and nitrous oxide at Station ALOHA, Limnol. Oceanogr., 41, 1619&amp;ndash;1628, 1996. </reference>
		<reference numeration="22" content_type="text"> Dore, J. E., Brum, J. R., Tupas, L. M., and Karl, D. M.: Seasonal and interannual variability in sources of nitrogen supporting export in the oligotrophic subtropical North Pacific Ocean, Limnol. Oceanogr., 47(6), 1595&amp;ndash;1607, 2002. </reference>
		<reference numeration="23" content_type="text"> Dugdale, R. C. and Goering, J. J.: Uptake of new and regenerated forms of nitrogen in primary productivity, Limnol. Oceanogr., 12, 196&amp;ndash;206, 1967. </reference>
		<reference numeration="24" content_type="text"> Dugdale, R. W. and Wilkerson, F. P.: The use of $^15$N to measure nitrogen uptake in eutrophic oceans: experimental considerations, Limnol. Oceanogr., 31, 673&amp;ndash;689, 1986. </reference>
		<reference numeration="25" content_type="text"> Dugdale, R. W., Wilkerson, F. P., Chavez, F. P., and Barber, R. T.: Estimating new production in the equatorial Pacific at 150&amp;deg;W, J. Geophys. Res., 97, 681&amp;ndash;686, 1992. </reference>
		<reference numeration="26" content_type="text"> Dugdale, R. C. and Goering, J. J.: Uptake of new and regenerated forms of nitrogen in primary productivity, Limnol. Oceanogr., 12, 196&amp;ndash;206, 1967. </reference>
		<reference numeration="27" content_type="text"> Dugdale, R. C. and Wilkerson, F. P.: The use of $^15$N to measure nitrogen uptake in eutrophic oceans, experimental considerations, Limnol. Oceanogr., 31(4), 673&amp;ndash;689, 1986. </reference>
		<reference numeration="28" content_type="text"> Dugdale, R. C., Wilkerson, F. P., Barber, R., and Chavez, F.: Estimating new production in the equatorial Pacific Ocean at 150&amp;deg;W, J. Geophys. Res., 97(C1), 681&amp;ndash;686, 1992. </reference>
		<reference numeration="29" content_type="text"> Eppley, R. W. and Peterson, B. J.: Particulate organic matter flux and planktonic new production in the deep ocean, Nature, 282, 677&amp;ndash;680, 1979. </reference>
		<reference numeration="30" content_type="text"> Falcon, L. I., Carpenter, E. J., Cipriano, F., Bergman, B., and Capone, D. G.: N2 fixation by unicellular bacterioplankton from the Atlantic and pacific oceans: phylogeny and in situ rates, Appl. Environ. Microbiol., 765&amp;ndash;770, 2004. </reference>
		<reference numeration="31" content_type="text"> Fernandez, C. and Raimbault, P.:. Nitrogen regeneration in the NE Atlantic Ocean and its impact on seasonal new, regenerated and export production, Mar. Ecol. Progr. Ser., 337, 79&amp;ndash;92, 2007. </reference>
		<reference numeration="32" content_type="text"> Galloway, J. N., Schlesinger, W. H., Levy, H., Michaels, A., and Schnoor, J. L.: Nitrogen fixation: Anthropogenic enhancement-environmental response, Global Biogeochem. Cy., 9, 235&amp;ndash;252, 1995. </reference>
		<reference numeration="33" content_type="text"> Garcia, N., Raimbault, P., Gouze, E., and Sandroni, V.: Dinitrogen fixation and primary production in Western Mediterranean Sea, C. R. Biol., 329(9), 742&amp;ndash;750, 2006. </reference>
		<reference numeration="34" content_type="text"> Garcia, N., Raimbault, P., and Sandroni, V.: Seasonal nitrogen fixation and primary production in Southwest Pacific (New Caledonia): Some evidence for nanoplanktonic diazotrophy and rapid transfer of newly fixed nitrogen to picoplankton organisms, Mar. Ecol. Progr. Ser., 343, 25&amp;ndash;33, 2007. </reference>
		<reference numeration="35" content_type="text"> Gentilhomme, V. and Raimbault, P.: Absorption et régénération de l&apos;azote dans a zone frontale du courant algérien (Méditerranée Occidentale): Réévaluation de la production nouvelle, Oceanol. Acta., 17, 555&amp;ndash;562, 1994. </reference>
		<reference numeration="36" content_type="text"> Glibert, P. M., Lipschultz, F., McCarthy, J. J., and Altabet, M. A.: Isotope dilution models of uptake and remineralization of ammonium by marine plankton, Limnol. Oceanogr., 27, 639&amp;ndash;650, 1982. </reference>
		<reference numeration="37" content_type="text"> Grob, C., Ulloa, O., Claustre, H., Huot, Y., Alarcón, G., and Marie, D.: Contribution of picoplankton to the total particulate organic carbon concentration in the eastern South Pacific, Biogeosciences, 4, 837&amp;ndash;852, 2007. </reference>
		<reference numeration="38" content_type="text"> Grüber, N. and Sarmiento, J.: Global patterns of marine nitrogen fixation and denitrification, Global Biogeochem. Cy., 11, 235&amp;ndash;266, 1997. </reference>
		<reference numeration="39" content_type="text"> Hansell, D. A. Olson, D. B., Dentener, F., and Zamora, L. M.: Assessment of excess nitrate development in the subtropical North Atlantic, Mar. Chem., 106, 562&amp;ndash;579, doi:10.1016/j.marchem, 2007. </reference>
		<reference numeration="40" content_type="text"> Harrison, W. G., Platt, T., and Lewis, M. R.: f-ratio and its relationship to ambient nitrate concentration in coastal waters, J. Plankton Res., 9(1), 235&amp;ndash;248, 1987. </reference>
		<reference numeration="41" content_type="text"> Harrison, W. G., Harris, L. R., and Irwin, B. D.: The kinetic of nitrogen utilization in the oceanic mixed layer: nitrate and ammonium interactions at nanomolar concentrations, Limnol. Oceanogr., 41, 16&amp;ndash;32, 1996. </reference>
		<reference numeration="42" content_type="text"> Holmes, M. R., Aminot, A., Kerouel, R., Hooker, B. A., and Peterson, J. B.: A simple and precise method for measuring ammonium in marine and freshwater ecosystems, Can. J. Fish Aquat. Sci., 56, 1801&amp;ndash;1808, 1999. </reference>
		<reference numeration="43" content_type="text"> Karl, D. M.: Nutrients dynamics in the deep blue sea, Trends Microbiol. 10, 410&amp;ndash;418, 2002. </reference>
		<reference numeration="44" content_type="text"> Karl, D. M., Michaels, A., Bergman, B., et al.: Dinitrogen fixation in the world&apos;s oceans, Biogeochemistry, 57/58, 47&amp;ndash;98, 2002. </reference>
		<reference numeration="45" content_type="text"> Karl, D. M., Letelier, R., Tupas, L., Dore, J., and Christian, J. D. H.: The role of nitrogen fixation in biochemical cycling in the subtropical North Pacific Ocean, Nature, 386, 533&amp;ndash;538, 1997. </reference>
		<reference numeration="46" content_type="text"> Laws, E.: Isotope dilution models and the mistery of the vanishing 15N, limnol. Oceanogr., 29, 379&amp;ndash;386, 1984. </reference>
		<reference numeration="47" content_type="text"> Libby, P. S. and Wheeler, P. A.: Particulate and dissolved organic nitrogen in the central and eastern Equatorial Pacifi, Deep-Sea Res., 44, 345&amp;ndash;361, 1996. </reference>
		<reference numeration="48" content_type="text"> Lomas, M. W. and Lipschultz, F.: Forming the primary nitrite maximum: nitrifiers or phytoplankton?, Limnol. Oceanogr., 51, 2453&amp;ndash;2467, 2006. </reference>
		<reference numeration="49" content_type="text"> Mahaffey, C., Michaels, A. F., and Capone, D. G.: The conundrum of marine nitrogen fixation, Am. J. Sci., 305, 546&amp;ndash;595, 2005. </reference>
		<reference numeration="50" content_type="text"> Mahowald, N.: Dust sources and deposition during the last glacial maximum and current climate: A comparison of model results with paleodata from ice cores and marine sediments, J. Geophys. Res.-Atmos., 104, 15 895&amp;ndash;15 916, 1999. </reference>
		<reference numeration="51" content_type="text"> Maita, Y. and Yanada, M.: Vertical distribution of total dissolved nitrogen and dissolved organic nitrogen in seawater, Geochem. J., 24, 245&amp;ndash;254, 1990. </reference>
		<reference numeration="52" content_type="text"> Masquelier, S. and Vaulot, D.: Distribution of micro-organisms along a transect in the South-East Pacific Ocean (BIOSOPE cruise) from epifluorescence microscopy, Biogeosciences Discuss., 4, 2667&amp;ndash;2697, 2007. </reference>
		<reference numeration="53" content_type="text"> Menzel, D. W. and Ryther J. H.: The annual cycle of primary production in the Sargasso Sea off Bermuda, Deep Sea Res., 6, 351&amp;ndash;367, 1960. </reference>
		<reference numeration="54" content_type="text"> Menzel, D. W. and Ryther, J. H.: The composition of particulate organic matter in the western north Atlantic, Limnol. Oceanogr., 9, 179&amp;ndash;186, 1964. </reference>
		<reference numeration="55" content_type="text"> Michaels, A. F., Karl, D. M., and Capone, D. G.: Element stoichiometry, new production and nitrogen fixation, Oceanography, 14, 68&amp;ndash;77, 2001. </reference>
		<reference numeration="56" content_type="text"> Montoya, J. P., Voss, M., Kähler, P., and Capone, D. G.: A simple, high precision, high sensitivity tracer assay for N&lt;sub&gt;2&lt;/sub&gt; fixation, Appl. Environ. Microb., 62, 986&amp;ndash;993, 1996. </reference>
		<reference numeration="57" content_type="text"> Montoya, J. P., Holl, C. M., Zehr, J. P., Hansen, A., Villareal, T. A., and Capone, D. G.: High rates of N2 fixation by unicellular diazotrophs in the oligotroph Pacific, Nature, 430, 1027&amp;ndash;1031, 2004. </reference>
		<reference numeration="58" content_type="text"> Morel, A. and Maritorena, S.: Bio-optical properties of oceanic waters: a reappraisal, J. Geophys. Res., 106(C4), 7163&amp;ndash;7180, 2001. </reference>
		<reference numeration="59" content_type="text"> Morel, A., Gentili, B., Claustre, H., Babin, M., Bricaud, A., Ras, J., and Tieche, F.: Optical properties of the clearest natural waters, Limonl. Oceanogr., 52, 217&amp;ndash;229, 2007. </reference>
		<reference numeration="60" content_type="text"> Nixon, S. W.: Coastal eutrophication: a definition, social causes and future concerns, Ophelia, 41, 199&amp;ndash;220, 1995. </reference>
		<reference numeration="61" content_type="text"> Peltzer, E. T. and Hayward, N. A.: Spatial and temporal variability of total organic carbon along the 140&amp;deg; W in the equatorial Pacific ocean in 1992, Global Biogeochem. Cy., 6, 45&amp;ndash;76, 1996. </reference>
		<reference numeration="62" content_type="text"> Platt, T. and Harison, W. G.: Biogennic fluxes of carbon and oxygen in the ocean, Nature, 318, 55&amp;ndash;58, 1985. </reference>
		<reference numeration="63" content_type="text"> Priscu, J. C. and Downes, M. T.: Nitrogen uptake, ammonium oxidation and nitrous oxide (N&lt;sub&gt;2&lt;/sub&gt;O) levels in the coastal waters of western Cook Strait, New Zealand, Estuary. Coast Shel Sci., 20, 529&amp;ndash;542, 1985. </reference>
		<reference numeration="64" content_type="text"> Procter, L. M. and Fuhrman, J. A.: Viral mortality of marine bacteria and cyanobacteria, Nature, 343, 1327&amp;ndash;1332, 1990. </reference>
		<reference numeration="65" content_type="text"> Raimbault, P., Slawyk, G., Coste, B., and Fry, J.: Feasibility of using an automated colorimetric procedure for the determination of seawater nitrate in the 0 to 100 nM range: Examples from field and culture, Mar. Biol., 104, 347&amp;ndash;351, 1990. </reference>
		<reference numeration="66" content_type="text"> Raimbault, P., Garcia, N., and Cerutti, F.: Distribution of inorganic and organic nutrients in the South Pacific Ocean &amp;ndash; evidence for long-term accumulation of organic matter in nitrogen-depleted waters, Biogeosciences Discuss., 4, 3041&amp;ndash;3087, 2007. </reference>
		<reference numeration="67" content_type="text"> Raimbault, P., Slawyk, G., Boudjellal, B., Coatanoan, C., Conan, P., Coste, B., Garcia, N., Moutin, T., and Pujo-Pay, M.: Carbon and nitrogen uptake and export in the equatorial pacific at 150&amp;deg; W: Evidence of an efficient regenerated production cycle, J. Geophys. Res., 104(C2), 3341&amp;ndash;3356, 1999. </reference>
		<reference numeration="68" content_type="text"> Raimbault, P., Slawyk, G., and Garcia, N.: Comparison between chemical and isotopic measurements of biological nitrate utilization: further evidence of low new production levels in the equatorial Pacific, Mar. Biol., 136, 1147&amp;ndash;1155, 2000. </reference>
		<reference numeration="69" content_type="text"> Ras, J., Claustre, H., and Uitz, J.: Spatial variability of phytoplankton pigment distributions in the Subtropical South Pacific Ocean: comparison between in situ and predicted data, Biogeosciences Discuss., 4, 3409&amp;ndash;3451, 2007. </reference>
		<reference numeration="70" content_type="text"> Redfield, A. C., Ketchum, B. H., and Richards, F. A: The influence of organisms on the composition of seawater, in: The Sea, edited by: Hill, M. N., Wiley, 26&amp;ndash;77, 1963. </reference>
		<reference numeration="71" content_type="text"> Rees, A. P., Joint, I., and Donald, D. M.: Early spring bloom phytoplankton-nutrient dynamics at the celtic Sea Shelf edge, Deep Sea Res., 46, 483&amp;ndash;510, 1999. </reference>
		<reference numeration="72" content_type="text"> Ryther, J. H. and Menzel, D. W.:. The seasonal and geographical range of primary production in the Western Sargasso sea, Deep Sea Res., 6, 235&amp;ndash;238, 1960. </reference>
		<reference numeration="73" content_type="text"> Sahlsten, E.: Nitrogenous nutrition in the euphotic zone of the central North pacific Gyre, Mar. Biol., 96, 433&amp;ndash;439, 1987. </reference>
		<reference numeration="74" content_type="text"> Slawyk, G. and Collos, Y.: $^13$C and $^15$N uptake by marine phytoplankton III. Interactions in euphotic zone profiles of stratified oceanic areas, Mar. Ecol. Prog. Ser. 19, 223&amp;ndash;231, 1984. </reference>
		<reference numeration="75" content_type="text"> Slawyk, G. and Raimbault, P.: Simple procedure for the simultaneous recovery of dissolved inorganic and organic nitrogen in $^15$N-tracer experiments and improving the isotopic mass balance, Mar. Ecol. Prog. Ser., 124, 289&amp;ndash;299, 1995. </reference>
		<reference numeration="76" content_type="text"> Slawyk, G., Raimbault, P., and Garcia, N.: Measuring gross uptake of $^15$ N labeled nitrogen by marine phytoplankton without particulate matter collection: Evidence of low $^15$N losses to the dissolved organic nitrogen pool, Limnol. Oceanogr., 43(7), 1734&amp;ndash;1739, 1998. </reference>
		<reference numeration="77" content_type="text"> Slawyk, G., Raimbault, P., and Garcia, N.: Use of $^15$N to measure dissolved organic nitrogen release by marine phytoplankton (reply to comment by Bronk and Ward), Limnol. Oceanogr., 45(8), 884&amp;ndash;1886, 2000. </reference>
		<reference numeration="78" content_type="text"> Tedetti, M., Sempéré, R., Vasilkov, A., Charrière, B., Nérini, D., Miller, W. L., Kawamura, K., and Raimbault, P.: High penetration of ultraviolet radiation in the South East Pacific waters, Geophys. Res. Lett., 34, L12610, doi: 10.29/2007/GL029823, 2007. </reference>
		<reference numeration="79" content_type="text"> Toggeweiler, J. R.: Is the downward dissolved organic matter (DOM) flux important in carbon export, in: Productivity of the ocean: Present and past, edited by: Berger, W. H., Smetacek, V. S., and Wefer, G., Wiley-Intersci., New-York, 65&amp;ndash;84, 1989. </reference>
		<reference numeration="80" content_type="text"> Tréguer, P. and LeCorre, P.: Manuel d&apos;analyses des sels nutritifs dans l&apos;eau de mer: Utilisation de l&apos;Autoanalyser II Technicon, Univ. of Bretagne Occidentale, Lab. de Chim. Mar., Brest, France, 1975. </reference>
		<reference numeration="81" content_type="text"> Ward, B. B.: Nitrification and the marine nitrogen cycle, in: Microbial ecology of the oceans, edited by: Kirchman, D., Wiley-Liss Inc., 427&amp;ndash;454, 2000. </reference>
		<reference numeration="82" content_type="text"> Ward, B. B., Talbot, M. C., and Perry, M. J.: Contributions of phytoplankton and nitrifying bacteria to ammonium and nitrite dynamics in coastal waters, Cont. Shelf. Res., 3, 383&amp;ndash;398, 1984. </reference>
		<reference numeration="83" content_type="text"> Ward, B. B., Kilpatrick, K. A., Renger, E. H., and Eppley, R. W.: Biological nitrogen cycling in the nitracline, Limnol. Oceanogr., 34(3), 493&amp;ndash;513, 1989. </reference>
		<reference numeration="84" content_type="text"> Ward, B. B. and Zafiriou, O. C.: Nitrification and nitric oxide in the oxygen minimum of the eastern tropical North Pacific, Deep Sea Res., 35(7), 1127&amp;ndash;1442, 1988. </reference>
		<reference numeration="85" content_type="text"> Weiss, R. F.: The solubility of nitrogen, oxygen and argon in water and seawater, Deep Sea-Res., 17, 721&amp;ndash;735, 1970. </reference>
		<reference numeration="86" content_type="text"> Zehr, J. P., Mellon, M. T., and Zani, S.: New nitrogen fixing microorganisms detected in oligotrophic oceans by the amplification of nitrogenase (\textitnifH) genes, Appl. Environ. Microbiol., 64, 3444&amp;ndash;3450, 1998. </reference>
		<reference numeration="87" content_type="text"> Zehr, J. P, Carpenter, E. J., and Villareal, T. A.: New perspectives on nitrogen-fixing microorganisms in tropical and subtropical oceans, Trends Microbiol., 8, 68&amp;ndash;73, 2000. </reference>
		<reference numeration="88" content_type="text"> Zehr, J. P. and Ward, B. B.: Nitrogen cycling in the Ocean: New perspectives on processes and paradigms, Appl. Environ. Microbiol., 68(3), 1015&amp;ndash;1024, 2002. </reference>
		<reference numeration="89" content_type="text"> Zehr, J. P., Waterbury, J. P., Turner, P. J., Montoya, J. P., Omoregie, E., Steward, G. F, Hansen, A., and Karl, D. M.: Unicellular cyanobacteria fix N2 in the subtropical North Pacific Ocean, Nature, 412 635&amp;ndash;638, 2001. </reference>
	</references>
</article>

