<?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-5-847-2008</doi>
	<article_url>http://www.biogeosciences.net/5/847/2008/</article_url>
	<abstract_html>http://www.biogeosciences.net/5/847/2008/bg-5-847-2008.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/5/847/2008/bg-5-847-2008.pdf</fulltext_pdf>
	<start_page>847</start_page>
	<end_page>864</end_page>
	<publication_date>2008-05-19</publication_date>
	<article_title content_type="html">Climate-mediated changes to mixed-layer properties in the Southern Ocean: assessing the phytoplankton response</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>P. W. Boyd</name>
			<email>pboyd@alkali.otago.ac.nz</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>S. C. Doney</name>
		</author>
		<author numeration="3" affiliations="1,3">
			<name>R. Strzepek</name>
		</author>
		<author numeration="4" affiliations="2">
			<name>J. Dusenberry</name>
		</author>
		<author numeration="5" affiliations="4">
			<name>K. Lindsay</name>
		</author>
		<author numeration="6" affiliations="5">
			<name>I. Fung</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">NIWA Centre for Chemical and Physical Oceanography, Dept. of Chemistry, University of Otago, Dunedin, New Zealand</affiliation>
		<affiliation numeration="2" content_type="html">Marine Chemistry and Geochemistry Department, Woods Hole Oceanographic Institution, Woods Hole, MA 02543, USA</affiliation>
		<affiliation numeration="3" content_type="html">Department of Chemistry, University of Otago, Dunedin, New Zealand</affiliation>
		<affiliation numeration="4" content_type="html">Climate and Global Dynamics, National Center for Atmospheric Research Boulder, CO 80307, USA</affiliation>
		<affiliation numeration="5" content_type="html">Department of Earth &amp; Planetary Science, Berkeley Institute of the Environment, Berkeley, CA 94720-1250, USA</affiliation>
	</affiliations>
	<abstract content_type="html">Concurrent changes in ocean chemical and physical properties influence
phytoplankton dynamics via alterations in carbonate chemistry, nutrient and
trace metal inventories and upper ocean light environment. Using a fully
coupled, global carbon-climate model (Climate System Model 1.4-carbon), we
quantify anthropogenic climate change relative to the background natural
interannual variability for the Southern Ocean over the period 2000 and
2100. Model results are interpreted using our understanding of the
environmental control of phytoplankton growth rates &amp;ndash; leading to two major
findings. Firstly, comparison with results from phytoplankton perturbation
experiments, in which environmental properties have been altered for key
species (e.g., bloom formers), indicates that the predicted rates of change
in oceanic properties over the next few decades are too subtle to be
represented experimentally at present. Secondly, the rate of secular climate
change will not exceed background natural variability, on seasonal to
interannual time-scales, for at least several decades &amp;ndash; which may not
provide the prevailing conditions of change, i.e. constancy, needed for
phytoplankton adaptation. Taken together, the relatively subtle
environmental changes, due to climate change, may result in adaptation by
resident phytoplankton, but not for several decades due to the confounding
effects of climate variability. This presents major challenges for the
detection and attribution of climate change effects on Southern Ocean
phytoplankton. We advocate the development of multi-faceted tests/metrics
that will reflect the relative plasticity of different phytoplankton
functional groups and/or species to respond to changing ocean conditions.</abstract>
	<references>
		<reference numeration="1" content_type="text">Alvaina, S., Moulin, C., Dandonneau, Y., Loisel, H. Bre&apos;on, F.-M.: A species-dependent bio-optical model of case I waters for global ocean color processing, Deep-Sea Res. I, 53, 917&amp;ndash;925, 2006. </reference>
		<reference numeration="2" content_type="text">Arrigo, K. R., Robinson, D. H., Worthen, D. L., Dunbar, R. B., DiTullio, G. R., VanWoert, M., and Lizotte, M. P.: Phytoplankton community structure and the drawdown of nutrients and CO&lt;sub&gt;2&lt;/sub&gt; in the Southern Ocean, Science, 283, 365&amp;ndash;367, 1999. </reference>
		<reference numeration="3" content_type="text">Baker, A. C., Starger, C. J., McClanahan, T. R., and Glynn, P. W.: Corals&apos; adaptive response to climate change, Nature, 430, 741, 2004. </reference>
		<reference numeration="4" content_type="text">Beardall, J. and Giordano, M.: Ecological implications of microalgal and cyanobacterial CO&lt;sub&gt;2&lt;/sub&gt; concentrating mechanisms, and their regulation, Funct. Plant Biol., 29, 335&amp;ndash;347, 2002. </reference>
		<reference numeration="5" content_type="text">Beninca, E., Huisman, J., Heerkloss, R., D. Jo\&quot; hn, K.D., Branco, P., Van Nes, E.H., Scheffer, M., and Ellner, S. P.: Chaos in a long-term experiment with a plankton community, Nature, 451, 822&amp;ndash;828, doi:10.1038/nature06512, 2008. </reference>
		<reference numeration="6" content_type="text">Bindoff, N. L., Willebrand, J., Artale, V., Cazenave, A., Gregory, J., Gulev, S., Hanawa, K., LeQuéré, C., Levitus, S., Nojiri, Y., Shum, C. K., Talley L. D., and Unnikrishnan A.: Observations: Oceanic Climate Change and Sea Level, in: Climate Change 2007: The Physical Science Basis, Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K. B., Tignor, M. and Miller, H. L., Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA, 2007. </reference>
		<reference numeration="7" content_type="text">Blackmon, M., Boville, B., Bryan, F., Dickinson, R., Gent,P., Kiehl, K., Moritz, R., Randall, D., Shukla, J., Solomon, S., Bonan, G., Doney, S., Fung, I., Hack, J., Hunke, E., Hurrell, J., Kutzbach, J., Meehl, J., Otto-Bliesner, B., Saravanan, R., Schneider, E. K., Sloan, L., Spall, M., Taylor, K., Tribbia J., and Washington, W.: The Community Climate System Model, Bull. Amer. Meteorol. Soc., 82, 2357&amp;ndash;2376, 2001. </reference>
		<reference numeration="8" content_type="text">Bopp, L., Monfray, P., Aumont, O., Dufresne, J.-L., Le Treut, H., Madec, G., Terray L., and Orr, J. C.: Potential impact of climate change on marine export production, Global Biogeochem. Cy., 15, 81&amp;ndash;99, 2001. </reference>
		<reference numeration="9" content_type="text">Boyd, P. W.: Environmental factors controlling phytoplankton processes in the Southern Ocean, J. Phycol, 38, 844&amp;ndash;861, 2002. </reference>
		<reference numeration="10" content_type="text">Boyd, P. W. and Doney, S. C.: Modelling regional responses by marine pelagic ecosystems to global climate change, Geophys. Res. Lett., 29(16), 53-1 to 53-4, doi:10.1029/2001GL014130, 2002. </reference>
		<reference numeration="11" content_type="text">Boyd, P. and Doney, S. C.: The impact of climate change and feedback process on the ocean carbon cycle, in: Ocean Biogeochemistry, edited by: Fasham, M., Springer, 157&amp;ndash;193, 2003. </reference>
		<reference numeration="12" content_type="text">Boyd, P. W., Law, C. S., Hutchins, D. A., Abraham, E. R., Croot, P. L., Ellwood, M., Frew, R.D., Hadfield, M., Hall, J., Handy, S., Hare, C., Higgins, J., Hill, P., Hunter, K. A., LeBlanc, K., Maldonado, M. T., McKay, R. M., Mioni, C., Oliver, M., Pickmere, S., Pinkerton, M., Safi, K., Sander, S., Sanudo-Wilhelmy, S. A., Smith, M., Strzepek, R., Tovar-Sanchez, A., and Wilhelm, S. W.: FeCycle: Attempting an iron biogeochemical budget from a mesoscale SF$_6$ tracer experiment in unperturbed low iron waters, Global Biogeochem. Cy., 19, GB4S20, doi:10.1029/2005GB002494, 2005. </reference>
		<reference numeration="13" content_type="text">Boyd, P. W., Jickells, T., Law, C. S., Blain, S., Boyle, E. A., Buesseler, K. O., Coale, K. H., Cullen, J. J., de Baar, H. J. W., Follows, M., Harvey, M., Lancelot, C., Levasseur, M., Owens, N. P. J., Pollard, R., Rivkin, R. B., Sarmiento, J., Schoemann, V. Smetacek, V., Takeda, S., Tsuda, A., Turner, S., and Watson, A. J.: Mesoscale iron enrichment experiments 1993&amp;ndash;2005: Synthesis and future directions, Science, 315, 612&amp;ndash;617, 2007. </reference>
		<reference numeration="14" content_type="text">Carpenter, S. R. and Cottingham, K. L.: Resilience and restoration of lakes, Conservation Ecology (online): available from the Internet: http://www.consecol.org/vol1/iss1/art2/, 1997. </reference>
		<reference numeration="15" content_type="text">Ciotti, A. M., Cullen, J. J. and M. R. Lewis, M. R.: Assessment of the relationships between dominant cell size in natural phytoplankton communities and the spectral shape of the absorption coefficient, Limnol. Oceanogr., 47, 404&amp;ndash;417, 2002. </reference>
		<reference numeration="16" content_type="text">Clarke, A.: Costs and consequences of evolutionary temperature adaptation, Trends Ecol. Evol., 18, 573&amp;ndash;581, 2003. </reference>
		<reference numeration="17" content_type="text">Cloern, J. E., Grenz, C., and Vidergar-Lucas, L.: An empirical model of the phytoplankton chlorophyll : carbon ratio-the conversion factor between productivity and growth rate, Limnol. Oceanogr., 40, 1313&amp;ndash;1321, 1995. </reference>
		<reference numeration="18" content_type="text">Collins, S. and Bell, G.: Phenotypic consequences of 1000 generations of selection at elevated CO&lt;sub&gt;2&lt;/sub&gt; in a green alga, Nature, 431, 566&amp;ndash;569, 2004. </reference>
		<reference numeration="19" content_type="text">Cottrell, M. T. and Suttle, C. A.: Production of Axenic Cultures of Micromonas-Pusilla (Prasinophyceae) Using Antibiotics, J. Phycol., 29, 385&amp;ndash;387, 1993. </reference>
		<reference numeration="20" content_type="text">Cubillos, J. C., Wright, S. W., Nash, G., de Salas, M. F., Griffiths, B., Tilbrook, B., Poisson, A., and Hallegraeff, G. M.: Shifts in geographic distributions of calcification morphotypes of the coccolithophorid Emiliania huxleyi in the Southern Ocean during 2001&amp;ndash;2006, Mar. Ecol. Prog. Ser., 348, 47&amp;ndash;58, 2007. </reference>
		<reference numeration="21" 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="22" content_type="text">Denman, K., Hofmann, E., and Marchant, H.: Marine biotic responses and feedbacks to environmental change and feedbacks to climate, in: Climate Change 1995, edited by: Houghton, J. T., Meira Filho, L. G., Callander, B. A., et al., The Science of Climate Change, Cambridge University Press, 1996. </reference>
		<reference numeration="23" content_type="text">Denman, K. L., Brasseur, G., Chidthaisong, A., Ciais, P., Cox, P. M., Dickinson, R. E.,Hauglustaine, D., Heinze, C., Holland, E., Jacob, D., Lohmann, U., Ramachandran, S., da Silva Dias, P. L., Wofsy S. C., and Zhang, X.: Couplings Between Changes in the Climate System and Biogeochemistry, in: Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K. B., Tignor, M., and Mille, H. L., Cambridge University Press, Cambridge, UK, 2007. </reference>
		<reference numeration="24" content_type="text">Dilling, L., Doney, S. C., Edmonds, J., Gurney, K. R., Harriss, R., Schimel, D., Stephens, B., and Stokes, G.: The role of carbon cycle observations and knowledge in carbon management, Ann. Rev. Env. Resour., 28, 521&amp;ndash;558, doi:10.1146/annurev.energy.28.011503.163443, 2003. </reference>
		<reference numeration="25" content_type="text">Doney, S. C., Bullister, J. L., and Wanninkhof, R.: Climatic variability in upper ocean ventilation diagnosed using chlorofluorocarbons, Geophys. Res. Lett., 25, 1399&amp;ndash;1402, 1998. </reference>
		<reference numeration="26" content_type="text">Doney, S. C., Lindsay, K., Caldeira, K., Campin, J.-M., Drange, H., Dutay, J.-C., Follows, M., Gao, Y., Gnanadesikan, A., Gruber, N., Ishida, A., Joos, F., Madec, G., Maier-Reimer, E., Marshall, J. C., Matear, R. J., Monfray, P., Mouchet, A., Najjar, R., Orr, J. C., Plattner, G.- K., Sarmiento, J., Schlitzer, R., Slater, R., Totterdell, I. J., Weirig, M.-F., Yamanaka, Y., and Yool, A.: Evaluating global ocean carbon models: the importance of realistic physics, Global Biogeochem. Cy., 18, GB3017, doi:10.1029/2003GB002150, 2004. </reference>
		<reference numeration="27" content_type="text">Doney, S. C., Lindsay, K., Fung I., and John, J.: Natural variability in a stable 1000 year coupled climate-carbon cycle simulation, J. Climate, 19, 3033&amp;ndash;3054, 2006. </reference>
		<reference numeration="28" content_type="text">Doney, S. C., Yeager, S., Danabasoglu, G., Large, W. G., and McWilliams, J. C.: Mechanisms governing interannual variability of upper ocean temperature in a global hindcast simulation, J. Phys. Oceanogr., 37, 1918&amp;ndash;1938, 2007. </reference>
		<reference numeration="29" content_type="text">Duce, R. A. and Tindale, N. W.: The atmospheric transport of iron and its deposition in the ocean, Limnol. Oceanogr., 36, 1715&amp;ndash;1726, 1991. </reference>
		<reference numeration="30" content_type="text">Dunne, J. A., Saleska, S. R., Fischer, M. L., and Harte, J.: Integrating experimental and gradient methods in ecological climate change research, Ecology, 85 904&amp;ndash;916, 2004. </reference>
		<reference numeration="31" content_type="text">Edwards, R. and Sedwick, P.: Iron in East Antarctic snow: Implications for atmospheric iron deposition and algal production in Antarctic waters. Geophys. Res. Lett., 28, 3907&amp;ndash;3910, 2001. </reference>
		<reference numeration="32" content_type="text">Falkowski, P. G. and LaRoche, J.: Acclimation to spectral irradiance in algae, J. Phycol., 27, 8&amp;ndash;14, 1991. </reference>
		<reference numeration="33" content_type="text">Falkowski P. G., Barber, R. T., and V. Smetacek, V.: Biogeochemical controls and feedbacks on ocean primary production, Science, 281, 200&amp;ndash;206, 1998. </reference>
		<reference numeration="34" content_type="text">Falkowski, P. G., Katz, M. E., Knoll, A. H., Quigg, A., Raven, J. A., Schofield, O., and Taylor, F. J. R.: The Evolution of Modern Eukaryotic Phytoplankton, Science 305, 354, doi:10.1126/science.1095964, 2004. </reference>
		<reference numeration="35" content_type="text">Feller, G., Ellis-Evans, J. C., Deubert, C., and Connelly, D. P.: Regulation by low temperature of phytoplankton growth and nutrient uptake in the Southern ocean, Mar. Ecol. Prog. Ser., 219, 51&amp;ndash;64, 2001. </reference>
		<reference numeration="36" content_type="text">Follows, M. J., Dutkiewicz S., Grant, S., and Chisholm, S. W.: Emergent Biogeography of Microbial Communities in a Model Ocean, Science, 1843&amp;ndash;1846, 2007. </reference>
		<reference numeration="37" content_type="text">Friedlingstein, P., Cox, P., Betts, R., Bopp, L., von Bloh, W., Brovkin, V., Cadule, S. Doney, M. Eby, I. Fung, G. Bala, J. John, C. Jones, F. Joos, T. Kato, P., Kawamiya, M., Knorr, W., Lindsay, K., Matthews, H. D., Raddatz, T., Rayner, P., Reick, C., Roeckner, E., Schnitzler, K.-G., Schnur, R., Strassmann, K., Weaver, A. J., Yoshikawa, C., and Zeng, N.: Climate&amp;ndash;carbon cycle feedback analysis: Results from the C4MIP model intercomparison, J. Climate, 19(14), 3337&amp;ndash;3353, 2006. </reference>
		<reference numeration="38" content_type="text">Fung, I. Y., Meyn, S. K., Tegen, I., Doney, S. C., John, J. G., and Bishop, J. K. B.: Iron supply and demand in the upper ocean, Global Biogeochem. Cy., 14, 697&amp;ndash;700, 2000. </reference>
		<reference numeration="39" content_type="text">Fung, I., Doney, S. C., Lindsay, K., and John, J.: Evolution of carbon sinks in a changing climate, Proc. Nat. Acad. Sci. (USA), 102, 11 201&amp;ndash;11 206, doi:10.1073/pnas.050494910, 2005. </reference>
		<reference numeration="40" content_type="text">Gibbs, S.J., Bown, P.R., Sessa, J.A., Bralower, T.J. and Wison, P.A.: Nannoplankton extinction and origination across the Paleocene-Eocene Thermal Maximum, Sceince, 314, 1770&amp;ndash;1773, doi:10.1126/science.1133902, 2006. </reference>
		<reference numeration="41" content_type="text">Gille, S. T.: Warming of the Southern Ocean since the 1950s. Science, 295, 5558, 1275&amp;ndash;1277, 2002. </reference>
		<reference numeration="42" content_type="text">Gordon, L. I., Codispoti, L. A., Jennings, J. C., Millero, F. J., Morrison, J. M., and Sweeney, C.: Seasonal evolution of hydrographic properties in the Ross Sea, Antarctica, 1996&amp;ndash;1997, Deep Sea Res. II, 47, 3095&amp;ndash;311, 2000. </reference>
		<reference numeration="43" content_type="text">Harrison, P. J., Waters, R. E., and Taylor, F. J. R.: A broad spectrum artificial seawater medium for coastal and open ocean phytoplankton, J. Phycol., 16, 28&amp;ndash;35, 1980. </reference>
		<reference numeration="44" content_type="text">Hiscock, M. R., Marra, J., Smith, W. O., Barber, R. T.: Primary productivity and its regulation in the Pacific Sector of the Southern Ocean, Deep Sea Res. II, 50, 533&amp;ndash;558, 2001. </reference>
		<reference numeration="45" content_type="text">Hutchins, D., Sedwick, P., DiTullio, G., Boyd, P., Quéguiner, B., Griffiths, B.F., and Crossley, C.: Control of phytoplankton growth by iron and silicic acid availability in the subantarctic Southern Ocean: Experimental results from the SAZ Project, J. Geophys. Res., 106, 31 559&amp;ndash;31 572, 2001. </reference>
		<reference numeration="46" content_type="text">Iglesias-Rodriguez, M. D., Brown, C. W., Doney, S. C., Kleypas, J., Kolber, D., Kolber, Z., Hayes, P. K., and Falkowski, P. G.: Representing key phytoplankton functional groups in ocean carbon cycle models: coccolithophores, Global Biogeochem. Cycles, 16, 1100, doi:10.1029/2001GB001454, 2002. </reference>
		<reference numeration="47" content_type="text">Iglesias-Rodriguez, M. D., Schofield, O. M., Batley, J., Medlin L. K., and Hayes, P. K.: Intraspecific genetic diversity in the marine coccolithophore Emiliania huxleyi (Prymnesiophyceae): The use of microsatellite analysis in marine phytoplankton population studies, J. Phycol. 42, 526&amp;ndash;536, 2006. </reference>
		<reference numeration="48" content_type="text">Iglesias-Rodriguez, M. D., Halloran, P. R., Rickaby, R. E. M., Hall, I. R., Colmenero-Hidalgo, E., Gittins, J. R., Green, D. R. H., Tyrrell, T., Gibbs, S. J., von Dassow, P., Rehm, E., Armbrust, E. V., and Boessenkool, K. P.: Phytoplankton Calcification in a High-CO&lt;sub&gt;2&lt;/sub&gt; World, Science 320, 336, doi: 10.1126/science.1154122, 2008. </reference>
		<reference numeration="49" content_type="text">Kaplan, A., Schwarz, R., Lieman-Hurwitz, J., and Reinhold L.: Physiological and Molecular Aspects of the Inorganic Carbon-Concentrating Mechanism in Cyanobacteria, Plant Physiol., 97, 851&amp;ndash;855, 1991. </reference>
		<reference numeration="50" content_type="text">Kepner, R. L. and Pratt, J. R.: Use of Fluorochromes for Direct Enumeration of Total Bacteria in Environmental-Samples &amp;ndash; Past and Present, Microbiol. Rev. 58, 603&amp;ndash;15, 1994. </reference>
		<reference numeration="51" content_type="text">Langer G., Geisen, M., Baumann, K.-H, Klaus, J., Riebesell, U., Thoms, S., and Young, J. R.: The response of Calcidiscus leptoporus and Coccolithus pelagicus to changing carbonate chemistry of seawater, Geophys. Res. Abst., 8, 05161, 2006a. </reference>
		<reference numeration="52" content_type="text">Langer G., Geisen M., Baumann K.-H., Klaus, J., Riebesell, U., Thoms, S., and Young, J. R.: Species-specific responses of calcifying algae to changing seawater carbonate chemistry, Geochem. Geophys. Geosyst., 7, Q09006, doi:10.1029/2005GC001227, 2006b. </reference>
		<reference numeration="53" content_type="text">Legendre L. and Rivkin, R. B.: Integrating functional diversity, food web processes, and biogeochemical carbon fluxes into a conceptual approach for modeling the upper ocean in a high-CO&lt;sub&gt;2&lt;/sub&gt; world, J. Geophys. Res., 110, C09S17, doi:10.1029/2004JC002530, 2005. </reference>
		<reference numeration="54" content_type="text">Le Quéré C., Harrison, S. P., Prentice, C., Buitenhuis, E. T., Aumont, O., Bopp, L., Claustre, H., Cotrim Da Cunha, L., Geider, R., Giraud, X., Klaas, C., Kohfeld, K., Legendre, L., Manizza, M., Platt, T., Rivkin, R. B., Sathyendranath, S., Uitz, J., Watson, A. J., and Wolf-Gladrow, D.: Ecosystem dynamics based on phytoplankton functional types for global ocean biogeochemistry models, Global Change Biol. 11, 2016&amp;ndash;2040, 2005. </reference>
		<reference numeration="55" content_type="text">Le Quéré, C., R\&quot; odenbeck, C., Buitenhuis, E. T. Conway, T. J. Langenfelds, R. Gomez, A. Labuschagne, C. Ramonet, M. Nakazawa, T. Metzl N., Gillett, N. and Heimann M.: Saturation of the Southern Ocean CO&lt;sub&gt;2&lt;/sub&gt; Sink Due to Recent Climate Change, Science, 316, 1735&amp;ndash;1738, doi:10.1126/science.1136188, 2007. </reference>
		<reference numeration="56" content_type="text">Levitus, S., Antonov, J. I., and Boyer, T. P: Warming of the World Ocean, 1955&amp;ndash;2003, Geophys. Res. Lett., 32, L02604, doi:10.1029/2004GL021592, 2005. </reference>
		<reference numeration="57" content_type="text">Litchman, E., Klausmeier, C. A., Miller, J. R., Schofield, O. M., and Falkowski, P. G.: Multinutrient, multi-group model of present and future oceanic phytoplankton communities, Biogeosciences, 3, 585&amp;ndash;606, http://www.biogeosciences.net/3/585/2006/. 2006, </reference>
		<reference numeration="58" content_type="text">Lovenduski, N. S., Gruber, N., Doney S. C., and Lima, I. D.: Enhanced CO&lt;sub&gt;2&lt;/sub&gt; outgassing in the Southern Ocean from a positive phase of the Southern Annular Mode, Global Biogeochem. Cy., 21, GB2026, doi:10.1029/2006GB002900, 2007. </reference>
		<reference numeration="59" content_type="text">MacIntyre, H. L. and Cullen, J. J.: Using cultures to investigate the Physiological ecology of Microalgae, in: Algal Culturing Techniques, edited by: Andersen, R. A., Elsevier, Amsterdam, 287&amp;ndash;326, 2005. </reference>
		<reference numeration="60" content_type="text">Mahowald, N. M. and Luo, C.: A less dusty future?, Geophys. Res. Lett., 30(17), 1903, doi:10.1029/2003GL017880, 2003. </reference>
		<reference numeration="61" content_type="text">Mahowald, N. M., Baker, A. R., Bergametti, G., Brooks, N., Duce, R. A., Jickells, T. D., Kubilay, N., Prospero, J. M., and Tegen, I.: Atmospheric global dust cycle and iron inputs to the ocean, Global Biogeochem. Cy., 19, GB4025, doi:10.1029/2004GB002402, 2005. </reference>
		<reference numeration="62" content_type="text">Maldonado, M. T. and Price, N. M.:. Influence of N substrate on Fe requirements of marine centric diatoms, Mar. Ecol. Prog. Ser, 141, 161&amp;ndash;172, 1996. </reference>
		<reference numeration="63" content_type="text">Maldonado, M. T. and Price, N. M.: Utilization of iron bound to strong organic ligands by plankton communities in the subarctic Pacific Ocean. Deep-Sea Res. II, 46, 2447&amp;ndash;2473, 1999. </reference>
		<reference numeration="64" content_type="text">Margalef, R: Life-forms of phytoplankton as survival alternatives in an unstable environment, Oceanologia Acta, 1, 493&amp;ndash;509, 1978. </reference>
		<reference numeration="65" content_type="text">Martin, C. L. and Tortell, P. D.: Bicarbonate transport and extracellular carbonic anhydrase activity in Bering Sea phytoplankton assemblages: Results from isotope disequilibrium experiments, Limnol. Oceanogr. 51, 2111&amp;ndash;2121, 2006. </reference>
		<reference numeration="66" content_type="text">Matear, R. and Hirst, A. C.: Climate change feedback on the future oceanic CO&lt;sub&gt;2&lt;/sub&gt; uptake. Tellus 51, 722&amp;ndash;733, 1999. </reference>
		<reference numeration="67" content_type="text">Measures C. I. and Vink, S.: Dissolved Fe in the upper waters of the Pacific sector of the Southern Ocean, Deep Sea Res. II, 48, 3913&amp;ndash;3941, 2001. </reference>
		<reference numeration="68" content_type="text">Medlin, L. K.: Genetic Variability and phytoplankton species, Chapter 34, IOC Workshop report #, 142, 257&amp;ndash;265, 1994. </reference>
		<reference numeration="69" content_type="text">Medlin, L. K., Barker, G. L. A., Wrieden, S., and Vaulot, D.: Genetic characterisation of Emiliania huxleyi (Haptophyta), J. Mar. Syst., 9, 13&amp;ndash;31, 1996. </reference>
		<reference numeration="70" content_type="text">Merico, A., Tyrrell, T., Brown, C. W., Groom, S. B., and Miller, P. I.: Analysis of satellite imagery for Emiliania huxleyi blooms in the Bering Sea before 1997, Geophys. Res. Lett., 30, 1337&amp;ndash;1340, 2003. </reference>
		<reference numeration="71" content_type="text">Moline, M. A., Claustre, H., Frazer, T. K., Schofield, O., and Vernet, M.: Alteration of the food web along the Antarctic Peninsula in response to a regional warming trend, Global Change Biol., 10, 1973&amp;ndash;1980, doi:10.1111/j.1365-2486.2004.00825, 2004. </reference>
		<reference numeration="72" content_type="text">Moran, X. A. G., Sebastián, M., Pedros-Alio, C., and Estrada, M.: Response of Southern ocean phytoplankton and bacterioplankton production to short-term experimental warming, Limnol. Oceanogr., 51, 1791&amp;ndash;1800, 2006. </reference>
		<reference numeration="73" content_type="text">Najjar, R. G., Jin, X., Louanchi, F., Aumont, O., Caldeira, K., Doney, S. C., Dutay, J.-C., Follows, M., Gruber, N., Joos, F., Lindsay, K., Maier-Reimer, E., Matear, R. J., Matsumoto, K., Monfray, P., Mouchet, A., Orr, J. C., Plattner, G. K., Sarmiento, J. L., Schlitzer, R., Slater, R.D., Weirig, M.-F., Yamanaka, Y., and Yool, A.: Impact of circulation on export production, dissolved organic matter and dissolved oxygen in the ocean: Results from Phase II of the Ocean Carbon-cycle Model Intercomparison Project (OCMIP-2), Global Biogeochem. Cy., 21, GB3007, doi:10.1029/2006GB002857, 2007. </reference>
		<reference numeration="74" content_type="text">Nelson, D. M., Brzezinski, M. A., Sigmon, D. E., and Franck, V. M.: A seasonal progression of Si limitation in the Pacific sector of the Southern Ocean, Deep-Sea Res. II, 48, 3973&amp;ndash;3995, 2001. </reference>
		<reference numeration="75" content_type="text">Nodder, S. D., Boyd, P. W., Chiswell, S. M., Pinkerton, M. H., Bradford-Grieve, J. M., Greig, M. J. N.: Temporal coupling between surface and deep ocean biogeochemical processes in contrasting subtropical and subantarctic water masses, southwest Pacific Ocean, J. Geophys. Res., 110, C12017, doi:10.1029/2004JC002833, 2005. </reference>
		<reference numeration="76" content_type="text">Peterson, W. T. and Keister, J. E.: Interannual variability in copepod community composition at a coastal station in the northern California Current: A multivariate approach, Deep-Sea Res. II, 50, 2499&amp;ndash;2517, 2003. </reference>
		<reference numeration="77" content_type="text">Price, N. M., Harrison, G. I., Hering, J. G., Hudson, R. J., Nirel, P. M. V., Palenik, B., and Morel, F. M. M.: Preparation and chemistry of the artificial algal culture medium Aquil, Biol. Oceanogr., 6, 443&amp;ndash;461, 1988. </reference>
		<reference numeration="78" content_type="text">Quigg, A., Finkel, Z. V., Milligan, A. J., Wyman, K., Falkowski, P. G., and Morel, F. M. M.: The evolutionary inheritance of elemental stoichiometry in marine phytoplankton, Nature, 425, 291&amp;ndash;294, 2003. </reference>
		<reference numeration="79" content_type="text">Raitsos, D. E., Lavender, S. J., Maravelias, C.D., Haralambous, J., Richardson, A.J., and P.C. Reid, P.C.: Identifying phytoplankton functional groups from space: an ecological approach. Limnol. Oceanogr., in press, 2008. </reference>
		<reference numeration="80" content_type="text">Raven, J. A., Evans, M. C. W., and Korb, R. E.: The role of trace metals in photosynthetic electron transport in O&lt;sub&gt;2&lt;/sub&gt;-evolving organisms, Photosynth. Res., 60, 111&amp;ndash;149, 1999. </reference>
		<reference numeration="81" content_type="text">Reynolds, C. S.: The Ecology of Freshwater Phytoplankton, 397 pp., Cambridge University Press, Cambridge, UK, 1984. </reference>
		<reference numeration="82" content_type="text">Riebesell, U., Zondervan, I., Rost, B., Tortell, P. D., Zeebe, R. E., and Morel, F. M. M.: Reduced calcification of marine plankton in response to increased atmospheric CO&lt;sub&gt;2&lt;/sub&gt;, Nature, 407, 364&amp;ndash;367, 2000. </reference>
		<reference numeration="83" content_type="text">Riebesell, U., Schulz, K. G., R. G. J. Bellerby, R. G. J., Botros, M., Fritsche, P., Meyerho\&quot; fer, M.,. Neill, N., Nondal, G., Oschlies, A., Wohlers, J., and Zöllner., E: Enhanced biological carbon consumption in a high CO&lt;sub&gt;2&lt;/sub&gt; ocean, Nature, 450, 545&amp;ndash;546, doi:10.1038/nature06267, 2007. </reference>
		<reference numeration="84" content_type="text">Ringbom, A.: Complexation in Analytical Chemistry, New York: Interscience, 1963. </reference>
		<reference numeration="85" content_type="text">Rost, B., Riebesell, U., and Burkhardt, S.: Carbon acquisition of bloom-forming marine phytoplankton, Limnol. Oceanogr., 48, 55&amp;ndash;67, 2003 </reference>
		<reference numeration="86" content_type="text">Russell, J. L., Dixon, K. W., Gnanadesikan, A., Stouffer, R. J., and Toggweiler, J. R.: The Southern Hemisphere westerlies in a warming world: propping open the door to the deep ocean, J. Climate, 19, 6382&amp;ndash;6390, 2006. </reference>
		<reference numeration="87" content_type="text">Saito, M. A., Sigman, D. M., and Morel, F. M. M: The bioinorganic chemistry of the ancient ocean: the co-evolution of cyanobacterial metal requirements and biogeochemical cycles at the Archean-Proterozoic boundary?, Inorg. Chim. Acta, 356, 308&amp;ndash;318, 2003. </reference>
		<reference numeration="88" content_type="text">Sarmiento, J. L.: Atmospheric CO&lt;sub&gt;2&lt;/sub&gt; stalled, Nature, 365, 697&amp;ndash;698, 1993. </reference>
		<reference numeration="89" content_type="text">Sarmiento, J. L., Hughes, T. M. C., Stouffer, R. J., and Manabe, S.: Simulated response of the ocean carbon cycle to anthropogenic climate warming, Nature, 393, 245&amp;ndash;249, 1998. </reference>
		<reference numeration="90" content_type="text">Sarmiento, J. L., Slater, R. Barber, R., Bopp, L., Doney, S. C., Hirst, A. C., Kleypas, J., Matear, R., Mikolajewicz, U., Monfray, P., Soldatov, V., Spall, S. A., and Stouffe, R.: Response of ocean ecosystems to climate warming, Global Biogeochem. Cy., 18, GB3003, doi:10.1029/2003GB002134, 2004. </reference>
		<reference numeration="91" content_type="text">Sedwick, P. N., Garcia, N. S., Riseman, S. F., Marsay, C. M., and DiTullio, G. R.: Evidence for high iron requirements of colonial Phaeocystis antarctica at low irradiance, Biogeochemistry, 83, 83&amp;ndash;97, doi:10.1007/s10533-007-9081-7, 2007. </reference>
		<reference numeration="92" content_type="text">Sherlock, V., Pickmere, S., Currie, K., Hadfield, M., Nodder, S., and Boyd, P. W.: Predictive accuracy of temperature-nitrate relationships for the oceanic mixed layer of the New Zealand region, J. Geophys. Res., 112, C06010, doi:10.1029/2006JC003562, 2007. </reference>
		<reference numeration="93" content_type="text">Smayda, T. J., Borkman, D. G., Beaugrand, G., and Belgrano, A. G.: Ecological effects of climate variation in the North Atlantic: Phytoplankton, in: Ecological Effects of Climatic Variations in the North Atlantic, edited by: Stenseth, N. C., Ottersen, G., Hurrell, J., and Belgrano, A., Oxford Univ. Press., UK, 48&amp;ndash;58, 2005. </reference>
		<reference numeration="94" content_type="text">Smetacek, V. and Nichol, S.: Polar ocean ecosystems in a changing world, Nature, 437, 362&amp;ndash;368, 2005. </reference>
		<reference numeration="95" content_type="text">Smith, T. M. and Reynolds, R. W.: A global merged land and sea surface temperature reconstruction based on historical observations (1880&amp;ndash;1997), J. Climate, 18, 2021&amp;ndash;2036, 2005. </reference>
		<reference numeration="96" content_type="text">Takeda, S.: Influence of iron availability on nutrient consumption ratio of diatoms in oceanic waters, Nature, 393, 774&amp;ndash;777, 1998. </reference>
		<reference numeration="97" content_type="text">Tilzer, M. M., Elbrächter, M., Gieskes, W. W., and Beese, B.: Light-Temperature interactions in the control of photosynthesis in Antarctica phytoplankton, Polar Biol., 5, 105&amp;ndash;111, 1986. </reference>
		<reference numeration="98" content_type="text">Timmermans, K. R., Gerringa, L. J. A., de Baar, H. J. W., van der Wagt, B., Veldhuis, M. J. W., de Jong, J. T. M., and Croot, P. L.: Growth rates of large and small Southern ocean diatoms in relation to the availability of iron in natural seawater, Limnol. Oceanogr., 46, 260&amp;ndash;266, 2001. </reference>
		<reference numeration="99" 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. Progr. Ser., 236, 37&amp;ndash;43, 2002. </reference>
		<reference numeration="100" content_type="text">Tortell, P. D., Martin and C. L., and Corkum, M. E.: Inorganic carbon uptake and intracellular assimilation by subarctic Pacific phytoplankton assemblages, Limnol. Oceanogr. 51, 2102&amp;ndash;2110, 2006. </reference>
		<reference numeration="101" content_type="text">Tortell, P. D., Payne, C. D., Li, Y., Trimborn, S., Rost, B., Smith, W. O., Riesselman, C., Dunbar R. B., Sedwick, P., and G. R. DiTullio, G.R.: ~CO&lt;sub&gt;2&lt;/sub&gt; sensitivity of Southern Ocean phytoplankton, Geophys. Res. Lett., 35, L04605, doi.org/10.1029/2007GL032583, 2008. </reference>
		<reference numeration="102" content_type="text">Tortell, P.D., Rost, B., Strzepek, R., Boyd, P.W., and DiTullio G.: Uptake and assimilation of inorganic carbon by Southern Ocean phytoplankton, Limnol. Oceanogr., in press, 2008. </reference>
		<reference numeration="103" content_type="text">Trenberth, K. E., Jones, P. D., Ambenje, P., Bojariu, R., Easterling, D., Klein, A., Tank, T., Parker, D., Rahimzadeh, F., Renwick, J. A., Rusticucci, M., Soden, B., and Zhai, P.: Observations: Surface and Atmospheric Climate Change, in: Climate Change 2007: The Physical Science Basis, Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K. B., Tignor, M., and Miller, H. L., Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA, 2007. </reference>
		<reference numeration="104" content_type="text">Walther, G.-R.: Tackling ecological complexity in climate impact research, Science, 315, 606&amp;ndash;607, 2007. </reference>
		<reference numeration="105" content_type="text">Westall, J. C., Zachary, J. L., and Morel, F. M. M.: MINEQL: A computer program for the calculation of chemical equilibrium composition of aqueous systems, (Department of Civil engineering, MIT, Cambridge, Massachusetts), 1976. </reference>
		<reference numeration="106" content_type="text">Westberry, T. K. and Siegel, D. A.: Spatial and temporal distribution of Trichodesmium blooms in the world&apos;s oceans, Global Biogeochem. Cy., 20, GB4016, doi:10.1029/2005GB002673, 2006. </reference>
	</references>
</article>

