<?xml version="1.0" encoding="utf-8" standalone="no"?>
<!DOCTYPE article SYSTEM "http://www.biogeosciences.net/inc/bg/copernicus.dtd">
<article language="en">
	<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-5-739-2008</doi>
	<article_url>http://www.biogeosciences.net/5/739/2008/</article_url>
	<abstract_html>http://www.biogeosciences.net/5/739/2008/bg-5-739-2008.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/5/739/2008/bg-5-739-2008.pdf</fulltext_pdf>
	<start_page>739</start_page>
	<end_page>748</end_page>
	<publication_date>2008-05-07</publication_date>
	<article_title content_type="html">Effects of increased atmospheric CO&lt;sub&gt;2&lt;/sub&gt; on small and intermediate sized osmotrophs during a nutrient induced phytoplankton bloom</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>A. I. Paulino</name>
		</author>
		<author numeration="2" affiliations="1">
			<name>J. K. Egge</name>
		</author>
		<author numeration="3" affiliations="1">
			<name>A. Larsen</name>
			<email>aud.larsen@bio.uib.no</email>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Department of Biology, Microbiology, University of Bergen, P. Box 7800, N-5020 Bergen, Norway</affiliation>
	</affiliations>
	<abstract content_type="html">We report the transient population dynamic response of the osmotrophic
community initiated by a nutrient pulse in mesocosms exposed to different
&lt;i&gt;p&lt;/i&gt;CO&lt;sub&gt;2&lt;/sub&gt; levels. Differences in phytoplankton and heterotrophic bacteria
abundances associated with the CO&lt;sub&gt;2&lt;/sub&gt; treatment are also described.
Coastal seawater was enclosed in floating mesocosms (27 m&lt;sup&gt;3&lt;/sup&gt;) and
nutrients were supplied initially in order to stimulate growth of microbial
organisms, including the coccolitophorid &lt;i&gt;Emiliania huxleyi&lt;/i&gt;. The mesocosms were modified to
achieve 350 μatm (1&amp;times;CO&lt;sub&gt;2&lt;/sub&gt;), 700 μatm (2&amp;times;CO&lt;sub&gt;2&lt;/sub&gt;) and 1050 μatm (3&amp;times;CO&lt;sub&gt;2&lt;/sub&gt;)
CO&lt;sub&gt;2&lt;/sub&gt; pressure. The temporal dynamics was related to
nutrient conditions in the enclosures. Numerically small osmotrophs
(picoeukaryotes and &lt;i&gt;Synechoccocus&lt;/i&gt; sp.) dominated initially and towards the end of the
experiment, whereas intermediate sized osmotrophs bloomed as the initial
bloom of small sized osmotrophs ceased. Maximum concentrations of &lt;i&gt;E. huxleyi&lt;/i&gt; were
approximately 4.6&amp;times;10&lt;sup&gt;3&lt;/sup&gt; cells ml&lt;sup&gt;&amp;minus;1&lt;/sup&gt; whereas other intermediate
sized osmotrophs reached approximately twice as high concentrations. The
osmotrophic succession pattern did not change, and neither were we able to
detect differences with regard to presence or absence of specific
osmotrophic taxa as a consequence of altered &lt;i&gt;p&lt;/i&gt;CO&lt;sub&gt;2&lt;/sub&gt;. Towards the end of
the experiment we did, however, record significantly higher picoeukaryotic-
and lower &lt;i&gt;Synechococcus&lt;/i&gt;-abundances in the higher CO&lt;sub&gt;2&lt;/sub&gt; treatments. Slightly increased
cell concentrations of &lt;i&gt;E. huxleyi&lt;/i&gt; and other nanoeukaryotes were also recorded at
elevated &lt;i&gt;p&lt;/i&gt;CO&lt;sub&gt;2&lt;/sub&gt; on certain days.</abstract>
	<references>
		<reference numeration="1" content_type="text">Badger, M. R. and Price, G. D.: CO&lt;sub&gt;2&lt;/sub&gt; concentrating mechanisms in cyanobacteria: molecular components, their diversity and evolution, J. Exp. Bot., 54, 609&amp;ndash;622, 2003. </reference>
		<reference numeration="2" content_type="text">Bratbak, G. and Thingstad, T. F.: Phytoplankton-bacteria interactions: an apparent paradox? Analysis of a model system with both competition and commensalism, Mar. Ecol.- Prog. Ser., 25, 23&amp;ndash;30, 1985. </reference>
		<reference numeration="3" content_type="text">Brewer, P. G., Goyet, C., and Friederich, G.: Direct observation of the oceanic CO&lt;sub&gt;2&lt;/sub&gt; increase revisited. P. Natl. Acad. Sci. USA, 94, 8308&amp;ndash;8313, 1997. </reference>
		<reference numeration="4" content_type="text">Brussaard, C. P. D.: Viral Control of Phytoplankton Populations &amp;ndash;- a Review, J. Eukaryot. Microbiol., 51, 125&amp;ndash;138, 2004. </reference>
		<reference numeration="5" content_type="text">Burkhardt, S., Riebesell, U., and Zondervan, I.: Effects of growth rate, CO&lt;sub&gt;2&lt;/sub&gt; concentration, and cell size on the stable carbon isotope fractionation in marine phytoplankton, Geochim. Cosmochim. Ac., 63, 3729&amp;ndash;3741, 1999. </reference>
		<reference numeration="6" content_type="text">Burkhardt, S., Amoroso, G., Riebesell, U., and Sultemeyer, D.: CO&lt;sub&gt;2&lt;/sub&gt; and HCO&lt;sub&gt;3&lt;/sub&gt; - uptake in marine diatoms acclimated to different CO&lt;sub&gt;2&lt;/sub&gt; concentrations, Limnol. Oceanogr., 46, 1378&amp;ndash;1391, 2001. </reference>
		<reference numeration="7" content_type="text">Cole, J. J., Findlay, S., and Pace, M. L.: Bacterial production in fresh and saltwater ecosystems: a cross-system overview, Mar. Ecol.- Prog. Ser., 43, 1&amp;ndash;10, 1988. </reference>
		<reference numeration="8" content_type="text">Delille, B., Harlay, J., Zondervan, I., Jacquet, S., Chou, L., Wollast, R., Bellerby, R. G. J., Frankignoulle, M., Borges, A. V., Riebesell, U., and Gattuso, J. P.: Response of primary production and calcification to changes of $p$CO&lt;sub&gt;2&lt;/sub&gt; during experimental blooms of the coccolithophorid Emiliania huxleyi, Global Biogeochem. Cy., 19, GB2023, doi:10.1029/2004GB002318,2005. </reference>
		<reference numeration="9" content_type="text">Egge, J. K.: Nutrient control of phytoplankton growth: Effects of macronutrient composition (N, P, Si) on species succession, Dr.s. thesis, University of Bergen, Norway. 40pp., 1993. </reference>
		<reference numeration="10" content_type="text">Egge, J. K., and Jacobsen, A.: Influence of silicate on particulate carbon production in phytoplankton, Mar. Ecol.-Prog. Ser., 147, 219&amp;ndash;230, 1997. </reference>
		<reference numeration="11" content_type="text">Egge, J. K., Thingstad T. F., Engel, A., and Riebesell, U.: Primary production during nutrient-induced blooms at elevated CO&lt;sub&gt;2&lt;/sub&gt; concentrations, Biogeosciences Discuss., 4, 3913&amp;ndash;3936, 2007. </reference>
		<reference numeration="12" content_type="text">Engel, A., Delille, B., Jacquet, S., Riebesell, U., Rochelle-Newall, E., Terbruggen, A., and Zondervan, I.: Transparent exopolymer particles and dissolved organic carbon production by Emiliania huxleyi exposed to different CO&lt;sub&gt;2&lt;/sub&gt; concentrations: A mesocosm experiment, Aquat. Microb. Ecol., 34, 93&amp;ndash;104, 2004. </reference>
		<reference numeration="13" content_type="text">Engel, A., Zondervan, I., Aerts, K., Beaufort, L., Benthien, A., Chou, L., Delille, B., Gattuso, J. P., Harlay, J., Heemann, C., Hoffmann, L., Jacquet, S., Nejstgaard, J., Pizay, M. D., Rochelle-Newall, E., Schneider, U., Terbrueggen, A., and Riebesell, U.: Testing the direct effect of CO&lt;sub&gt;2&lt;/sub&gt; concentration on a bloom of the coccolithophorid Emiliania huxleyi in mesocosm experiments, Limnol. Oceanogr., 50, 493&amp;ndash;507, 2005. </reference>
		<reference numeration="14" content_type="text">Engel, A., Schulz, K. G., Riebesell, U., Bellerby, R., Delille, B., Schartau, M.: Effects of CO&lt;sub&gt;2&lt;/sub&gt; on particle size distribution and phytoplankton abundance during a mesocosm bloom experiment (PeECE II), Biogeosciences, 5, 509&amp;ndash;521, 2008. </reference>
		<reference numeration="15" content_type="text">Fenchel, T.: Relation between particle size selection and clearance in suspension-feeding ciliates, Limnol. Oceanogr., 25, 733&amp;ndash;738, 1980. </reference>
		<reference numeration="16" content_type="text">Fenchel, T.: Ecology &amp;ndash; Potentials and Limitations, Excellence in Ecology Series, Vol. 1. Ecology Institute, 43&amp;ndash;54, 1987. </reference>
		<reference numeration="17" content_type="text">Fu, F. X., Warner, M. E., Zhang, Y. H., Feng, Y. Y., and Hutchins, D. A.: Effects of increased temperature and co2 on photosynthesis, growth, and elemental ratios in marine Synechococcus and Prochlorococcus (Cyanobacteria), J. Phycol., 43, 485&amp;ndash;496, 2007. </reference>
		<reference numeration="18" content_type="text">Giordano, M., Beardall, J., and Raven, J. A.: CO&lt;sub&gt;2&lt;/sub&gt; concentrating mechanisms in algae: Mechanisms, environmental modulations, and evolution, Annu. Rev. Plant. Biol., 56, 99&amp;ndash;131, 2005. </reference>
		<reference numeration="19" content_type="text">Grossart, H.P., and Simon, M.: Bacterial colonization and microbial decomposition of limnetic aggregates (lake snow), Aquat. Microb. Ecol., 15, 127&amp;ndash;140, 1998. </reference>
		<reference numeration="20" content_type="text">Grossart, H. P., Levold, F., Allgaier, M., Simon, M., and Brinkhoff, T.: Marine diatom species harbour distinct bacterial communities, Environ. Microbiol., 7, 860&amp;ndash;873, 2005. </reference>
		<reference numeration="21" content_type="text">Grossart, H. P., Allgaier, M., Passow, U., and Riebesell, U.: Testing the effect of CO&lt;sub&gt;2&lt;/sub&gt; concentration on the dynamics of marine heterotrophic bacterioplankton, Limnol Oceanogr, 51, 1&amp;ndash;11, 2006. </reference>
		<reference numeration="22" content_type="text">Hamm, C. E.: Architecture, ecology and biogeochemistry of phaeocystis colonies, J. Sea Res., 43, 307&amp;ndash;315, 2000. </reference>
		<reference numeration="23" content_type="text">Hamm, C. E. M., Merkel, R., Springer, O., Jurkojc, P., Maier, C., Prechtel, K., and Smetacek, V.: Architecture and material properties of diatom shells provide effective mechanical protection, Nature, 421, 841&amp;ndash;843, 2003. </reference>
		<reference numeration="24" content_type="text">Hare, C. E., Leblanc, K., DiTullio, G. R., Kudela, R. M., Zhang, Y., Lee, P. A., Riseman, S., and Hutchins, D. A.: Consequences of increased temperature and CO&lt;sub&gt;2&lt;/sub&gt; for phytoplankton community structure in the Bering Sea, Mar. Ecol.-Prog. Ser., 352, 9&amp;ndash;16, 2007. </reference>
		<reference numeration="25" content_type="text">Hassidim, M., Keren, N., Ohad, I., Reinhold, L., and Kaplan, A.: Acclimation of Synechococcus strain WH7803 to ambient CO&lt;sub&gt;2&lt;/sub&gt; concentration and to elevated light intensity, J. Phycol., 33, 811&amp;ndash;817, 1997. </reference>
		<reference numeration="26" content_type="text">John, D. E., Wang, Z. A., Liu, X., Byrne, R. H., Corredor, J. E., López, J. M., Cabrera, A., Bronk, D., Tabita, F.R., and Paul, J. H.: Phytoplankton carbon fixation gene (RuBisCO) transcripts and air-sea CO&lt;sub&gt;2&lt;/sub&gt; flux in the Mississippi River Plume, The ISME Journal, 1, 517&amp;ndash;531, 2007. </reference>
		<reference numeration="27" content_type="text">Kuenen, J. G., Boonstra, J., Scroder, H. G. J., and Veldkamp, H.: Competition for inorganic substrates among chemoorganotrophic and chemolithtrophic bacteria. Microbial Ecol., 3, 119&amp;ndash;130, 1977. </reference>
		<reference numeration="28" content_type="text">Kuenzler, E. J. and Perras, J. P.: Phosphatases of marine algae, Biol. Bull., 128, 271&amp;ndash;284, 1965. </reference>
		<reference numeration="29" content_type="text">Larsen, J. B., Larsen, A., Thyrhaug, R., Bratbak, G., and Sandaa, R.-A.: Response of marine viral populations to a nutrient induced phytoplankton bloom at different $p$CO&lt;sub&gt;2&lt;/sub&gt; levels, Biogeosciences, 5, 523&amp;ndash;533, 2008. </reference>
		<reference numeration="30" content_type="text">Marie, D., Brussaard, C. P. D., Partensky, F., and Vaulot, D.: Enumeration of phytoplankton, bacteria, viruses in marine samples, in: Current Protocols in Cytometry, edited by: Robinson, J. P., Darzynkiewicz, Z., Dean, P. N., Orfao, A., and 4 others, John Wiley &amp; Sons, Chichester, 11.11.1&amp;ndash;11.11.15, 1999. </reference>
		<reference numeration="31" content_type="text">Not, F., Massana, R., Latasa, M., Marie, D., Colson, C., Eikrem, W., Pedros-Alio, C., Vaulot, D., and Simon, N.: Late summer community composition and abundance of photosynthetic picoeukaryotes in Norwegian and Barents Seas, Limnol. Oceanogr., 50, 1677&amp;ndash;1686, 2005. </reference>
		<reference numeration="32" content_type="text">Pinhassi, J., Sala, M. M., Havskum, H., Peters, F., Guadayol, O., Malits, A., and Marrase, C. L.: Changes in bacterioplankton composition under different phytoplankton regimens, Appl. Environ. Microb., 70, 6753&amp;ndash;6766, 2004. </reference>
		<reference numeration="33" content_type="text">Pinhassi, J., Gomez-Consarnau, L., Alonso-Saez, L., Sala, M. M., Vidal, M., Pedros-Alio, C., and Gasol, J. M.: 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="34" content_type="text">Raven, J. and Waite, A.: The evolution of silification in diatoms: inescapable sinking and sinking as escape, New Phytol., 162, 45&amp;ndash;65, 2004. </reference>
		<reference numeration="35" content_type="text">Riebesell, U.: Effects of CO&lt;sub&gt;2&lt;/sub&gt; enrichment on marine phytoplankton, J. Oceanogr., 60, 719&amp;ndash;729, 2004. </reference>
		<reference numeration="36" content_type="text">Riebesell, U., Revill, A. T., Holdsworth, D. G., and Volkman, J. K.: The effects of varying CO$_2 $ concentration on lipid composition and carbon isotope fractionation in Emiliania huxleyi, Geochimica Et Cosmochimica Acta, 64, 4179&amp;ndash;4192, 2000. </reference>
		<reference numeration="37" content_type="text">Riebesell, U., Schultz, K. G., Bellerby, R. G. J., Botros, M., Fritsche, P., Meyerhöfer, M., Neill, C., Nondal, G., Oschlies, A., Wohlers, J., and Zöllner, E.: Enhanced biological carbon consumption in high CO&lt;sub&gt;2&lt;/sub&gt; ocean, Nature, 450, 545&amp;ndash;548, doi:10.1038/nature06267, 2007. </reference>
		<reference numeration="38" content_type="text">Riegman, R., Stolte, W., Noordeloos, A. A. M., and Slezak, D.: Nutrient uptake and alkaline phosphatase (EC 3:1:3:1) activity of Emiliania huxleyi (Prymnesiophyceae) during growth under N and P limitation in continuous cultures, J. Phycol., 36, 87&amp;ndash;96, 2000. </reference>
		<reference numeration="39" content_type="text">Rochelle-Newall, E., Delille, B., Frankignoulle, M.., Gattuso, J.-P., Jacquet, S., Riebesell, U., Terbruggen, A., and Zondervan, I.: Chromophoric dissolved organic matter in experimental mesocosms maintained under different $p$CO&lt;sub&gt;2&lt;/sub&gt; levels, Mar. Ecol.- Prog. Ser., 272, 25&amp;ndash;31, 2004. </reference>
		<reference numeration="40" content_type="text">Rost, B., Riebesell, U., Burkhardt, S., and Sultemeyer, D.: Carbon acquisition of bloom-forming marine phytoplankton, Limnol. Oceanogr., 48, 55&amp;ndash;67, 2003. </reference>
		<reference numeration="41" content_type="text">Sandaa, R.-A., Heldal, M., Castberg, T., Thyrhaug, R., and Bratbak., G.: Isolation and characterization of two marine viruses infecting Chrysochromulina ericina and Pyramimonas orientalis, Virology, 290, 272&amp;ndash;280, 2001. </reference>
		<reference numeration="42" content_type="text">Schulz, K. G. and Riebesell, U.: Build-up and decline of organic matter during PeECE III, Biogeosciences, 5, 707&amp;ndash;718, 2008. </reference>
		<reference numeration="43" content_type="text">Shapiro J.: Blue-green algae: Why they become dominant, Science, 179, 382&amp;ndash;384, 1973. </reference>
		<reference numeration="44" content_type="text">Smith, R. E. H. and Kalff, J.: Size-dependent phosphorus uptake kinetics and cell quota in phytoplankton, J. Phycol., 18, 275&amp;ndash;284, 1982. </reference>
		<reference numeration="45" content_type="text">Smith, D. C., Steward, G. F., Long, R. A., and Azam, F.: Bacterial mediation of carbon fluxes during a diatom bloom in a mesocosm, Deep-Sea Res. Pt Ii, 42, 75&amp;ndash;97, 1995. </reference>
		<reference numeration="46" content_type="text">Sokal, R.R. and Rohlf, F.J.: Biometry, the principles and practice of statistics in biological research 3rd Ed. 7th Printing. W. H. Freeman and Company, 887 pp., 2001. </reference>
		<reference numeration="47" content_type="text">Suttle, C. and Chan, A.: Viruses infecting the marine prymnesiophyte Chrysochromulina spp.: isolation, preliminary characterization and natural abundance. Mar. Ecol.- Prog. Ser., 118, 275&amp;ndash;282, 1995. </reference>
		<reference numeration="48" content_type="text">Tanaka, T., Thingstad, T. F., Løvdal, T., Grossart, H. P., Larsen A., Schultz, K., Riebesell, U.: Availability of phosphate for phytoplankton and bacteria and of glucose for bacteria at different $p$CO&lt;sub&gt;2&lt;/sub&gt; levels in a mesocosm study, Biogeosciences, 5, 669&amp;ndash;678, 2008. </reference>
		<reference numeration="49" content_type="text">Thingstad, T. F.: A theoretical approach to structuring mechanisms in the pelagic food web, Hydrobiologia, 363, 59&amp;ndash;72, 1998. </reference>
		<reference numeration="50" content_type="text">Thingstad, T. F.: Elements of a theory for the mechanisms controlling abundance, diversity, and biogeochemical role of lytic bacterial viruses in aquatic systems, Limnol. Oceanogr., 45, 1320&amp;ndash;1328, 2000. </reference>
		<reference numeration="51" content_type="text">Thingstad, T. F. and Lignell, R.: Theoretical models for the control of bacterial growth rate, abundance, diversity and carbon demand, Aquat. Microb. Ecol., 13, 19&amp;ndash;27, 1997. </reference>
		<reference numeration="52" content_type="text">Thingstad, T. F., Havskum, H., Zweifel, U. L., Berdalet, E., Sala, M.M., Peters, F., Alcaraz, M., Scharek, R., Perez, M., Jacquet, S., Flaten, G. A. F., Dolan, J.R., Marrasé, C., Rassoulzadegan, F., Hagstrøm, A., and Vaulot, D.: Ability of a &quot;minimum&quot; microbial food web model to reproduce response patterns observed in mesocosms manipulated with N and P, glucose, and Si, J. Marine Syst., 64, 15&amp;ndash;34, 2007. </reference>
		<reference numeration="53" content_type="text">Thingstad, T. F., Ovreas, L., Egge, J. K., Lovdal, T., and Heldal, M.: Use of non-limiting substrates to increase size; a generic strategy to simultaneously optimize uptake and minimize predation in pelagic osmotrophs?, Ecol. Lett., 8, 675&amp;ndash;682, 2005. </reference>
		<reference numeration="54" content_type="text">Throndsen J., Hasle G. R., and Tangen K.: Norsk Kystplankton Flora, Oslo Almater Forlag AS, Norway, 2003. </reference>
		<reference numeration="55" content_type="text">Tortell, P. D.: Evolutionary and ecological perspectives on carbon acquisition in phytoplankton, Limnol Oceanogr., 45, 744&amp;ndash;750, 2000. </reference>
		<reference numeration="56" content_type="text">Tortell, P. D., Rau G. H., and Morell, F. M., M.: Inorganic carbon acquisition in coastal pacific phytoplankton communities, Limol. Oceanogr., 45, 1485&amp;ndash;1500, 2000. </reference>
		<reference numeration="57" content_type="text">Tortell, P. D., DiTullio, G. R., Sigman, D. M., and Morel, F. M. M.: CO&lt;sub&gt;2&lt;/sub&gt; effects on taxonomic composition and nutrient utilization in an equatorial pacific phytoplankton assemblage, Mar. Ecol.&amp;ndash;Prog. Ser., 236, 37&amp;ndash;43, 2002. </reference>
		<reference numeration="58" content_type="text">Williams, T. G., and Turpin, D. H.: Photosynthesis knetics determine the outcome of competition for dissolved inorganic carbon by freshwater microalgae: implications for acidified lakes, Oecologia (Berlin), 73, 307&amp;ndash;311, 1987. </reference>
	</references>
</article>

