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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>3</issue_number>
		<publication_year>2008</publication_year>
	</journal>
	<doi>10.5194/bg-5-833-2008</doi>
	<article_url>http://www.biogeosciences.net/5/833/2008/</article_url>
	<abstract_html>http://www.biogeosciences.net/5/833/2008/bg-5-833-2008.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/5/833/2008/bg-5-833-2008.pdf</fulltext_pdf>
	<start_page>833</start_page>
	<end_page>845</end_page>
	<publication_date>2008-05-19</publication_date>
	<article_title content_type="html">Factors limiting heterotrophic bacterial production in the southern Pacific Ocean</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>F. Van Wambeke</name>
			<email>france.van-wambeke@univmed.fr</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>S. Bonnet</name>
		</author>
		<author numeration="3" affiliations="3">
			<name>T. Moutin</name>
		</author>
		<author numeration="4" affiliations="3">
			<name>P. Raimbault</name>
		</author>
		<author numeration="5" affiliations="4">
			<name>G. Alarcón</name>
		</author>
		<author numeration="6" affiliations="2">
			<name>C. Guieu</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Laboratoire de Microbiologie, Géochimie et Ecologie Marines (LMGEM), CNRS, UMR 6117, Université de la Méditerranée, Campus de Luminy &amp;ndash; Case 901, 13 288 Marseille cedex 9, France</affiliation>
		<affiliation numeration="2" content_type="html">CNRS, Laboratoire d&apos;océanographie de Villefranche, 06 230 Villefranche-sur-Mer, France; Université Pierre et Marie Curie - Paris6, Laboratoire d&apos;océanographie de Villefranche, 06 230 Villefranche-sur-Mer, France</affiliation>
		<affiliation numeration="3" content_type="html">Laboratoire d&apos;Océanographie et de Biogéochimie (LOB), CNRS, UMR 6535, Université de la Méditerranée, Campus de Luminy &amp;ndash; Case 901, 13 288 Marseille cedex 9, France</affiliation>
		<affiliation numeration="4" content_type="html">Department of Oceanography &amp; Center for Oceanographic Research in the eastern South Pacific, University of Concepcion, Casilia 160-C, Concepcion, Chile</affiliation>
	</affiliations>
	<abstract content_type="html">The role of potential factors limiting bacterial growth was investigated
along vertical and longitudinal gradients across the South Eastern Pacific
Gyre. The effects of glucose, nitrate, ammonium and phosphate additions on
heterotrophic bacterial production (using leucine technique) were studied in
parallel in unfiltered seawater samples incubated under natural daily
irradiance. The enrichments realized on the subsurface showed three types of
responses. From 141&amp;deg; W (Marquesas plateau) to approx 125&amp;deg; W, bacteria
were not bottom-up controlled, as confirmed by the huge potential of growth
in non-enriched seawater (median of enhancement factor&amp;times;39 in 24 h). Within
the Gyre (125&amp;deg; W&amp;ndash;95&amp;deg; W), nitrogen alone stimulated leucine
incorporation rates (median&amp;times;4.2), but rapidly labile carbon (glucose)
became a second limiting factor (median&amp;times;37) when the two elements were
added. Finally from the border of the gyre to the Chilean upwelling (95&amp;deg; W&amp;ndash;73&amp;deg; W),
labile carbon was the only factor stimulating heterotrophic
bacterial production. Interaction between phytoplankton and heterotrophic
bacterial communities and the direct versus indirect effect of iron and
macronutrients on bacterial production were also investigated in four
selected sites: two sites on the vicinity of the Marquesas plateau, the
centre of the gyre and the Eastern border of the gyre. Both phytoplankton
and heterotrophic bacteria were limited by availability of nitrogen within
the gyre, but not by iron. Iron limited phytoplankton at Marquesas plateau
and at the eastern border of the gyre. However 48 h enrichment experiments
were not sufficient to show any clear limitation of heterotrophic bacteria
within Marquesas plateau and showed a limitation of these organisms by
labile carbon in the eastern border of the Gyre.</abstract>
	<references>
		<reference numeration="1" content_type="text">Arrieta, J., Weinbauer, M., Lute, C., and Herndl, G.: Response of bacterioplankton to iron fertilization in the southern Ocean, Limnol. Oceanogr., 49, 799&amp;ndash;808, 2004. </reference>
		<reference numeration="2" content_type="text">Blain, S., Bonnet, S., and Guieu, C.: Dissolved iron distribution in the tropical and sub tropical South Eastern Pacific, Biogeosciences, 5, 269&amp;ndash;280, 2008. </reference>
		<reference numeration="3" content_type="text">Bonnet, S., Guieu, C., Bruyant, F., Prášil, O., Van Wambeke, F., Raimbault, P., Moutin, T., Grob, C., Gorbunov, M., Zehr, J. P., Masquelier, S., Garczarek, L., and Claustre, H.: Nutrient limitation of primary productivity in the Southeast Pacific (BIOSOPE cruise), Biogeosciences, 5, 215&amp;ndash;2225, 2008. </reference>
		<reference numeration="4" content_type="text">Bruland, K. W., Franks, R. P., Knauer, G. A. and Martin, J. H: Sampling and analytical methods for the determination of Cu, Cd. Zn, and Ni at the nanogram per liter level in sea water, Anal. Chim. Acta, 105, 234&amp;ndash;245, 1979. </reference>
		<reference numeration="5" content_type="text">Carlson, C. A. and Ducklow, H. W.: Growth of bacterioplankton and consumption of dissolved organic carbon in the Sargasso Sea, Aquat. Microb. Ecol., 10, 69&amp;ndash;85, 1996. </reference>
		<reference numeration="6" content_type="text">Cherrier, J., Bauer, J. E., and Druffel, E. R. M.: Utilization and turnover of labile dissolved organic matter by bacterial heterotrophs in eastern North Pacific surface waters, Mar. Ecol. Prog. Ser., 139, 267&amp;ndash;279, 1996. </reference>
		<reference numeration="7" content_type="text">Church, M., Hutchins, D., and Ducklow, H. W.: Limitation of bacterial growth by dissolved organic matter and iron in the Southern Ocean, Appl. Environ. Microbiol., 66, 455&amp;ndash;466, 2000. </reference>
		<reference numeration="8" content_type="text">Claustre, H., Sciandra, A., and Vaulot, D.: Introduction to the special section Bio-optical and biogeochemical conditions in the South East Pacific in late 2004: the BIOSOPE program, Biogeosciences, 5, 679&amp;ndash;691, 2008. </reference>
		<reference numeration="9" content_type="text">Cochlan, W. P.: The heterotrophic bacterial response during a mesoscale iron enrichment experiment (IronEx II) in the Eastern equatorial Pacific Ocean, Limnol. Oceanogr., 46, 428&amp;ndash;435, 2001. </reference>
		<reference numeration="10" content_type="text">Cotner, J., Ammermann, J., Peele, E., and Bentzen, E.: Phosphorus-limited bacterioplankton growth in the Sargasso Sea, Aquat. Microb. Ecol., 13, 141&amp;ndash;149, 1997. </reference>
		<reference numeration="11" content_type="text">Duhamel, S., Moutin, T., Van Wambeke, F., Van Mooy, B., Rimmelin, P., Raimbault, P., and Claustre, H.: Growth and specific P-uptake rates of bacterial and phytoplanktonic communities in the Southeast Pacific (BIOSOPE cruise), Biogeosciences, 4, 941&amp;ndash;956, 2007. </reference>
		<reference numeration="12" content_type="text">Donachie, S. P., Christian, J. R., and Karl, D. M.: Nutrient regulation of bacterial production and ectoenzyme activities in the subtropical North Pacific Ocean, Deep-Sea Res. II, 48, 1719&amp;ndash;1732, 2001. </reference>
		<reference numeration="13" content_type="text">Granéli, W., Carlsson, P., and Bertilsson, S.: Bacterial abundance, production and organic carbon limitation in the Southern Ocean (39&amp;ndash;62&amp;deg; S, 4&amp;ndash;14&amp;deg; E) during the austral summer 1997/1998, Deep-Sea Res. II, 51, 2569&amp;ndash;2582, 2004. </reference>
		<reference numeration="14" 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="15" content_type="text">Hutchins, D. A., DiTullio, G. R., Zhang, Y., and Bruland, K. W.: An Iron limitation mosaic in the California upwelling regime, Limnol. Oceanogr., 43, 1037&amp;ndash;1054, 1998. </reference>
		<reference numeration="16" content_type="text">Kirchman, D. L.: The uptake of inorganic nutrients by heterotrophic bacteria, Microb. Ecol., 28, 255&amp;ndash;271, 1994. </reference>
		<reference numeration="17" content_type="text">Kirchman, D. L.: Limitation of bacterial growth by dissolved organic matter in the subarctic Pacific, Mar. Ecol. Prog. Ser., 62, 47&amp;ndash;54, 1990. </reference>
		<reference numeration="18" content_type="text">Kirchman, D. L. and Rich, J.: Regulation of bacterial growth rates by dissolved organic carbon and temperature in the Equatorial Pacific Ocean, Microb. Ecol., 33, 11&amp;ndash;20, 1997. </reference>
		<reference numeration="19" content_type="text">Kirchman, D. L., Meon, M., Cottrell, M. T., Hutchins, D. A., Weeks, D., and Bruland, K. W.: Carbon versus iron limitation of bacterial growth in the California upwelling regime, Limnol. Oceanogr., 45, 1681&amp;ndash;1688, 2000. </reference>
		<reference numeration="20" content_type="text">Kirchman, D. L., Elifantz, H., Dittel, A., Malmstrom, R., and Cottrell, M.: Standing stocks and activity of Archaea and Bacteria in the western Arctic Ocean, Limnol. Oceanogr., 52, 495&amp;ndash;507, 2007. </reference>
		<reference numeration="21" content_type="text">Kuipers, B.: Diel periodicity of bacterioplankton in the euphotic zone of the subtropical Atlantic Ocean, Mar. Ecol. Prog. Ser., 201, 13&amp;ndash;25, 2000. </reference>
		<reference numeration="22" content_type="text">McAndrew, P., Bjorkman, K., Church, M., Morris, P., Jachowski, N., Williams, P. J. le. B., and Karl, D.: Metabolic response of oligotrophic plankton communities to deep water nutrient enrichment, Mar. Ecol. Prog. Ser., 332, 63&amp;ndash;75, 2007. </reference>
		<reference numeration="23" content_type="text">Moutin, T., Karl, D. M., Duhamel, S., Rimmelin, P., Raimbault, P., Van Mooy, B. A. S., and Claustre, H.: Phosphate availability and the ultimate control of new nitrogen input by nitrogen fixation in the tropical Pacific Ocean, Biogeosciences, 5, 95&amp;ndash;109, 2008. </reference>
		<reference numeration="24" content_type="text">Obernosterer, I., Catala, P., Lami, R., Caparros, J., Ras, J., Bricaud, A., Dupuy, C., Van Wambeke, F. and Lebaron, P.: Biochemical characteristics and bacterial community structure of the sea surface microlayer in the South Pacific Ocean, Biogeosciences, 5: 693&amp;ndash;705, 2008a. </reference>
		<reference numeration="25" content_type="text">Obernosterer, I., Christaki, U., Lefèvre, D., Catala, P., Van Wambeke, F., and Lebaron, P.: Rapid bacterial mineralization of organic carbon produced during a phytoplankton bloom induced by natural iron fertilization in the Southern Ocean, Deep-Sea Res. II, 55, 777&amp;ndash;789, 2008b. </reference>
		<reference numeration="26" content_type="text">Olivier, J. L., Barber, R. T., Smith, W. O., and Ducklow, H. W.: The heterotrophic bacterial response during the southern Ocean Iron Experiment (SOFeX), Limnol. Oceanogr., 49, 2129&amp;ndash;2140, 2004. </reference>
		<reference numeration="27" content_type="text">Pakulski, J. D., Coffin, R. B., Kelley, C. A., Holder, S. L., Downer, R., Aas, P., Lyons, M. M., and Jeffrey, W. H.: Iron stimulation of Antarctic bacteria. Nature, 383, 133&amp;ndash;134, 1996. </reference>
		<reference numeration="28" content_type="text">Pinhassi, J., Gómez-Consarnau, L., Alonso-Sáez, L., Sala, M., Vidal, M., Pedrós-Alió, C., and Gasol, J.: Seasonal changes in bacterioplankton nutrient limitation and their effects on bacterial community composition in the NW Mediterranean Sea, Aquat. Microb. Ecol., 44, 241&amp;ndash;252, 2006. </reference>
		<reference numeration="29" content_type="text">Pomeroy, L. R., Sheldon, J. E., Sheldon, W. M., and Peters, F.: Limits to growth and respiration of bacterioplankton in the Gulf of Mexico, Mar. Ecol. Prog. Ser., 117, 259&amp;ndash;268, 1995. </reference>
		<reference numeration="30" content_type="text">Raimbault, P., Garcia, N.: Evidence for efficient regenerated production and dinitrogen fixation in nitrogen-deficient waters of the South Pacific Ocean: impact on new and export production estimates, Biogeosciences, 5, 323&amp;ndash;338, 2008 </reference>
		<reference numeration="31" content_type="text">Raimbault, P., Garcia, N., and Cerutti, F.: Distribution of inorganic and organic nutrients in the South Pacific Ocean - Evidence for long-term accumulation of organic matter in nitrogen-depleted waters, Biogeosciences, 5, 281&amp;ndash;298, 2008. </reference>
		<reference numeration="32" 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, 5, 353&amp;ndash;369, 2008. </reference>
		<reference numeration="33" content_type="text">Sala, M. M., Peters, F., Gasol, J. M., Pedrós-Alió, C., Marrassé, C., and Vaqué, D.: Seasonal and spatial variations in the nutrient limitation of bacterioplankton growth in the northwestern Mediterranean, Aquat. Microb. Ecol., 27, 47&amp;ndash;56, 2002. </reference>
		<reference numeration="34" content_type="text">Shiah, F. K.: Diel cycles of heterotrophic bacterioplankton abundance and production in the ocean surface waters, Aquat. Microb. Ecol., 17, 239&amp;ndash;246, 1999. </reference>
		<reference numeration="35" content_type="text">Sempéré, S., Tedetti, M., Charrière, B., Panagiotopoulos, C. and Van Wambeke, F.: Molecular distribution and bacterial availability of dissolved sugars in the south East Pacific, Biogeosciences Discuss., 5, 725&amp;ndash;750, 2008. </reference>
		<reference numeration="36" 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:10.1029/2007GL029823, 2007. </reference>
		<reference numeration="37" content_type="text">Thingstad, T.: Control of bacterial growth in idealized food webs, in: Microbial Ecology of the Oceans, edited by: Kirchman, D. L., Wiley-Liss, New York, 239&amp;ndash;260, 2000. </reference>
		<reference numeration="38" content_type="text">Thingstad, T., Krom, M., Mantoura, F., Flaten, G., Groom, S., Herut, B., Kress, N., Law, C., Pasternak, A., Pitta, P., Psarra, S., Rassoulzadegan, F., Tanaka, T., Tselepides, A., Wassmann, P., Woodward, M., Riser, C., Zodiatis, G., and Zohary, T.: Nature of phosphorus limitation in the ultraoligotrophic eastern Mediterranean, Science, 309, 1068&amp;ndash;1071, 2005. </reference>
		<reference numeration="39" content_type="text">Tortell, P. D., Maldonaldo, M. T., Granger, J., and Price, N. M.: Marine bacteria and biogeochemical cycling of iron in the oceans, FEMS Microb. Ecol., 29, 1&amp;ndash;11, 1999. </reference>
		<reference numeration="40" content_type="text">Tortell, P. D., Maldonado, M. T., and Price, N. M.: The role of heterotrophic bacteria in iron-limited ecosystems. Nature, 383, 330&amp;ndash;332, 1996. </reference>
		<reference numeration="41" content_type="text">Torréton, J.-P., Talbot, V., and Garcia, N.: Nutrient stimulation of bacterioplankton growth in Tuamotu atoll lagoons, Aquat. Microb. Ecol., 21, 125&amp;ndash;137, 2000. </reference>
		<reference numeration="42" content_type="text">Van Wambeke, F., Christaki, U., Giannakourou, A., Moutin, T., and Souvemerzoglou, K.: Longitudinal and vertical trends of bacterial limitation by phosphorus and carbon in the Mediterranean Sea, Microb. Ecol., 43, 119&amp;ndash;133, 2002. </reference>
		<reference numeration="43" content_type="text">Van Wambeke, F., Obernosterer, I., Moutin, T., Duhamel, S., Ulloa, O., and Claustre, H.: Heterotrophic bacterial production in the South East Pacific: Longitudinal trends and coupling with primary production, Biogeosciences, 5, 157&amp;ndash;169, 2008a. </reference>
		<reference numeration="44" content_type="text">Van Wambeke, F., Tedetti, M., Duhamel, S., and Sempéré, R.: Diel variability of heterotrophic bacterial production and UV doses in the South East Pacific, Biogeosciences Discuss., 5, 435&amp;ndash;462, 2008b. </reference>
		<reference numeration="45" content_type="text">Zohary, T., and Robarts, R. D.: Experimental study of microbial P limitation in the eastern Mediterranean, Limnol. Oceanogr., 43, 387&amp;ndash;395, 1998. </reference>
	</references>
</article>

