<?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>7</volume_number>
		<issue_number>2</issue_number>
		<publication_year>2010</publication_year>
	</journal>
	<doi>10.5194/bg-7-657-2010</doi>
	<article_url>http://www.biogeosciences.net/7/657/2010/</article_url>
	<abstract_html>http://www.biogeosciences.net/7/657/2010/bg-7-657-2010.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/7/657/2010/bg-7-657-2010.pdf</fulltext_pdf>
	<start_page>657</start_page>
	<end_page>669</end_page>
	<publication_date>2010-02-16</publication_date>
	<article_title content_type="html">Effects of an iron-light co-limitation on the elemental composition (Si, C, N) of the marine diatoms &lt;I&gt;Thalassiosira oceanica&lt;/I&gt; and &lt;I&gt;Ditylum brightwellii&lt;/I&gt;</article_title>
	<authors>
		<author numeration="1" affiliations="1,2">
			<name>E. Bucciarelli</name>
			<email>eva.bucciarelli@univ-brest.fr</email>
		</author>
		<author numeration="2" affiliations="1,2">
			<name>P. Pondaven</name>
		</author>
		<author numeration="3" affiliations="1,2">
			<name>G. Sarthou</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Université Européenne de Bretagne, France</affiliation>
		<affiliation numeration="2" content_type="html">Université de Brest, CNRS, IRD, UMR 6539 LEMAR, IUEM; Technopôle Brest Iroise, Place Nicolas Copernic, 29280 Plouzané, France</affiliation>
	</affiliations>
	<abstract content_type="html">We examined the effect of iron (Fe) and Fe-light (Fe-L) co-limitation on
cellular silica (BSi), carbon (C) and nitrogen (N) in two marine diatoms,
the small oceanic diatom &lt;I&gt;Thalassiosira oceanica&lt;/I&gt; and the large coastal species &lt;I&gt;Ditylum brightwellii&lt;/I&gt;. We showed that C and
N per cell tend to decrease with increasing Fe limitation (i.e. decreasing
growth rate), both under high light (HL) and low light (LL). We observed an
increase (&lt;I&gt;T. oceanica&lt;/I&gt;, LL), no change (&lt;I&gt;T. oceanica&lt;/I&gt;, HL) and a decrease (&lt;I&gt;D. brightwellii&lt;/I&gt;, HL and LL) in BSi per
cell with increasing degree of limitation. The comparison with literature
data showed that the trend in C and N per cell for other Fe limited diatoms
was similar to ours. Interspecific differences in C and N quotas of Fe
limited diatoms observed in the literature seem thus to be mostly due to
variations in cell volume. On the contrary, there was no global trend in BSi
per cell or per cell volume, which suggests that other interspecific
differences than Fe-induced variations in cell volume influence the degree
of silicification. The relative variations in C:N, Si:C and Si:N versus the
relative variation in specific growth rate (i.e. μ:μ&lt;sub&gt;max&lt;/sub&gt;)
followed the same patterns for &lt;I&gt;T. oceanica&lt;/I&gt; and &lt;I&gt;D. brightwellii&lt;/I&gt;, whatever the irradiance level.
However, the variations of C:N under Fe limitation reported in the
literature for other diatoms are contrasted, which may thus be more related
to growth conditions than to interspecific differences. As observed in other
studies, Si:C and Si:N ratios increased by more than 2-fold between 100%
and 40% of μ&lt;sub&gt;max&lt;/sub&gt;. Under more severe limitation (HL and LL), we
observed for the first time a decrease in these ratios. These results may
have important biogeochemical implications on the understanding and the
modelling of the oceanic biogeochemical cycles, e.g. carbon and silica
export.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Allen, A. E., La Roche, J., Maheswari, U., Lommer, M., Schauer, N., Lopez, P. J., Finazzi, G., Fernie, A. R., and Bowler, C.: Whole-cell response of the pennate diatom \textitPhaeodactylum tricornutum to iron starvation, Proc. Natl. Acad. Sci., 105, 10438–10443, 2008. </reference>
		<reference numeration="2" content_type="text"> Aumont, O., Maier-Reimer, E., Blain, S., and Monfray, P.: An ecosystem model of the global ocean including Fe, Si, P co-limitations, Global Biogeochem. Cycles, 17(2), 1060, doi:10.1029/2001GB001745, 2003. </reference>
		<reference numeration="3" content_type="text"> Barber, R. T. and Hiscock, M. R.: A rising tide lifts all phytoplankton: growth response of other phytoplankton taxa in diatom-dominated blooms, Global Biogeochem. Cycles, 20, GB4S03, doi:10.1029/2006GB002726, 2006. </reference>
		<reference numeration="4" content_type="text"> Beucher, C., Tréguer, P., Corvaisier, R., Hapette, A. M., and Elskens, M.: Production and dissolution of biosilica, and changing microphytoplankton dominance in the Bay of Brest (France), Mar. Ecol. Progr. Ser., 267, 57–69, 2004. </reference>
		<reference numeration="5" content_type="text"> Boyd, P., LaRoche, J., Gall, M., Frew, R., and McKay, R. M. L.: Role of iron, light, and silicate in controlling algal biomass in subantarctic waters SE of New Zealand, J. Geophys. Res., 104, 13395–13408, 1999. </reference>
		<reference numeration="6" content_type="text"> Boyle, E.: Pumping iron makes thinner diatoms, Nature, 393, 733–734, 1998. </reference>
		<reference numeration="7" content_type="text"> Chisholm, S. W., Azam, F., and Eppley, R. W.: Silicic acid incorporation in marine diatoms on light:dark cycles: use as an essay for phased cell division, Limnol. Oceanogr., 23, 518–529, 1978. </reference>
		<reference numeration="8" content_type="text"> Claquin, P., Martin-Jézéquel, V., Kromkamp, J. C., Veldhuis, M. J. W., and Kraay, G. W.: Uncoupling of silicon compared with carbon and nitrogen metabolisms and the role of the cell cycle in continuous cultures of \textitThalassiosira pseudonana (Bacillariophyceae) under light, nitrogen, and phosphorus control, J. Phycol., 38, 922–930, 2002. </reference>
		<reference numeration="9" content_type="text"> Claquin, P. and Bucciarelli, E.: Cell cycle and silicification: Impact of an iron-light co-limitation on the marine diatom \textitThalassiosira oceanica, in preparation, 2010. </reference>
		<reference numeration="10" content_type="text"> de Baar, H. J. W. and La Roche, J.: Trace metals in the oceans: evolution, biology and global change, in: Marine science frontiers for Europe, edited by: Wefer, G., Lamy, F., and Mantoura, F., Springer-Verlag, Berlin Heidelberg New York Tokyo, 79–105, 2003. </reference>
		<reference numeration="11" content_type="text"> de Baar, H. J. W., Boyd, P., Coale, K. H., Landry, M. R., Tsuda, A., Assmy, P., Bakker, D. C. E., Bozec, Y., Barber, R. T., Brzezinski, M. A., Buesseler, K. O., Boyé, M., Croot, P. L., Gervais, F., Gorbunov, M. Y., Harrison, P. J., Hiscock, W. T., Laan, P., Lancelot, C., Law, C. S., Levasseur, M., Marchetti, A., Millero, F. J., Nishioka, J., Nojiri, Y., van Oijen, T., Riebesell, U., Rijkenberg, M. J. A., Saito, H., Takeda, S., Timmermans, K. R., Veldhuis, M. J. W., Waite, A. M., and Wong, C. S.: Synthesis of iron fertilization experiments: from the Iron Age in the Age of Enlightenment, J. Geophys. Res., 110, C09S16, doi:10.1029/2004JC002601, 2005. </reference>
		<reference numeration="12" content_type="text"> Dugdale, R. C., Wilkerson, F. P., and Minas, H. J.: The role of a silicate pump in driving new production, Deep Sea Res., 42, 697–719, 1995. </reference>
		<reference numeration="13" content_type="text"> Eppley, R. W. and Rogers, J. N.: Inorganic nitrogen assimilation of \textitDitylum brightwellii, a marine plankton diatom, J. Phycol., 6, 344–351, 1970. </reference>
		<reference numeration="14" content_type="text"> Falkowski, P., Scholes, R. J., Boyle, E., Canadell, J., Canfield, D., Elser, J., Gruber, N., Hibbard, K., Högberg, P., Linder, S., Mackenzie, F. T., Moore III, B., Pedersen, T., Rosenthal, Y., Seitzinger, S., Smetacek, V., and Steffen, W.: The global carbon cycle: a test of our knowledge of Earth as a system, Science, 290, 291–296, 2000. </reference>
		<reference numeration="15" content_type="text"> Falkowski, P. G., Barber, R. T., and Smetacek, V.: Biogeochemical controls and feedbacks on ocean primary production, Science, 281, 200–206, 1998. </reference>
		<reference numeration="16" content_type="text"> Fasham, M. J. R., Flynn, K. J., Pondaven, P., Anderson, T. R., and Boyd, P. W.: Development of a robust marine ecosystem model to predict the role of iron in biogeochemical cycles: A comparison of results for iron-replete and iron-limited areas, and the SOIREE iron-enrichment experiment, Deep Sea Res., 53, 333–366, 2006. </reference>
		<reference numeration="17" content_type="text"> Firme, G. F., Rue, E. L., Weeks, D. A., Bruland, K. W., and Hutchins, D. A.: Spatial and temporal variability in phytoplankton iron limitation along the California coast and consequences for Si, N,and C biogeochemistry, Global Biogeochem. Cycles, 17, 1016, doi:1010.1029/2001GB001824, 2003. </reference>
		<reference numeration="18" content_type="text"> Gallinari, M., Bucciarelli, E., Moriceau, B., and Ragueneau, O.: Dissolution properties of biogenic silica from diatoms grown under iron-replete and iron-limited conditions, in preparation, 2010. </reference>
		<reference numeration="19" content_type="text"> Geider, R. J. and La Roche, J.: Redfield revisited: variability of C:N:P in marine microalgae and its biochemical basis, Eur. J. Phycol., 37, 1–17, 2002. </reference>
		<reference numeration="20" content_type="text"> Goldman, J. C.: Inorganic carbon availability and the growth of large marine diatoms, Mar. Ecol. Prog. Ser., 180, 81–91, 1999. </reference>
		<reference numeration="21" content_type="text"> Hamm, C. E., 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, 841–843, 2003. </reference>
		<reference numeration="22" content_type="text"> Hoffmann, L. J., Peeken, I., and Lochte, K.: Effects of iron on the elemental stoichiometry during EIFEX and in the diatoms \textitFragilariopsis kerguelensis and \textitChaetoceros dichaeta, Biogeosciences, 4, 569–579, 2007. </reference>
		<reference numeration="23" content_type="text"> Hoffmann, L. J., Peeken, I., and Lochte, K.: Iron, silicate, and light co-limitation of three Southern Ocean diatom species, Polar Biol., 31, 1067–1080, 2008. </reference>
		<reference numeration="24" content_type="text"> Hopkinson, B. M. and Barbeau, K. A.: Interactive influences of iron and light limitation on phytoplankton at subsurface chlorophyll maxima in the eastern North Pacific, Limnol. Oceanogr., 53, 1303–1318, 2008. </reference>
		<reference numeration="25" content_type="text"> Hudson, R. J. M. and Morel, F. M. M.: Iron transport in marine phytoplankton: kinetics of cellular and medium coordination reactions, Limnol. Oceanogr., 35, 1002–1020, 1990. </reference>
		<reference numeration="26" content_type="text"> Hutchins, D. A. and Bruland, K. W.: Iron-limited diatom growth and Si:N uptake ratios in a coastal upwelling regime, Nature, 393, 561–564, 1998. </reference>
		<reference numeration="27" content_type="text"> IPCC: IPCC fourth assessment report: Synthesis report, http://www.ipcc.ch/pdf/assessment-report/ar4/syr/ar4_syr.pdf, 2007. </reference>
		<reference numeration="28" content_type="text"> Irigoien, X., Flynn, K. J., and Harris, R. P.: Phytoplankton blooms: a `loophole&apos; in microzooplankton grazing impact?, J. Plankton Res., 27, 313–321, 2005. </reference>
		<reference numeration="29" content_type="text"> Jensen, S. and Bathmann, U.: Algae viability within copepod faecal pellets: evidence from microscopic examinations, Mar. Ecol. Progr. Ser., 337, 145–153, 2007. </reference>
		<reference numeration="30" content_type="text"> Keller, M. D., Bellows, W. K., and Guillard, R. R. L.: Microwave treatment for sterilization of phytoplankton culture media, J. Exp. Mar. Biol. Ecol., 117, 279–283, 1988. </reference>
		<reference numeration="31" content_type="text"> Landry, M. R., Selph, K. E., Brown, S. L., Abbott, M. R., Measures, C. I., Vink, S., Allen, C. B., Calbet, A., Christensen, S., and Nolla, H.: Seasonal dynamics of phytoplankton in the Antarctic Polar Front region at 170&amp;deg; W, Deep Sea Res., 49, 1843–1865, 2002. </reference>
		<reference numeration="32" content_type="text"> La Roche, J., Murray, H., Orellana, M., and Newton, J.: Flavodoxin expression as an indicator of iron limitation in marine diatoms, J. Phycol., 31, 520–530, 1995. </reference>
		<reference numeration="33" 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, 161–172, 1996. </reference>
		<reference numeration="34" content_type="text"> Maldonado, M. T., Boyd, P. W., Harrison, P. J., and Price, N. M.: Co-limitation of phytoplankton growth by light and Fe during winter in the NE subarctic Pacific Ocean, Deep Sea Res. II, 46, 2475–2485, 1999. </reference>
		<reference numeration="35" content_type="text"> Maldonado, M. T. and Price, N. M.: Nitrate regulation of Fe reduction and transport by Fe-limited \textitThalassiosira oceanica, Limnol. Oceanogr., 45, 814–826, 2000. </reference>
		<reference numeration="36" content_type="text"> Marchetti, A. and Harrison, P. J.: Coupled changes in the cell morphology and the elemental (C, N , and Si) composition of the pennate diatom \textitPseudo-nitzschia due to iron deficiency Limnol. Oceanogr., 52, 2270–2284, 2007. </reference>
		<reference numeration="37" content_type="text"> Marchetti, A. and Cassar, N.: Diatom elemental composition and morphological changes in response to iron limitation: a brief review with potential paleoceanographic applications, Geobiology, 7, 419–431, 2009. </reference>
		<reference numeration="38" content_type="text"> Martin-Jézéquel, V., Hildebrand, M., and Brzezinski, M. A.: Silicon metabolism in diatoms: implications for growth, J. Phycol., 36, 1–20, 2000. </reference>
		<reference numeration="39" content_type="text"> Moore, C. M., Mills, M. M., Milne, A., Langlois, R., Achterberg, E. P., Lochte, K., Geider, R. J., and La Roche, J.: Iron limits primary productivity during spring bloom development in the central North Atlantic, Global Change Biol., 12, 626–634, 2006. </reference>
		<reference numeration="40" content_type="text"> Moore, C. M., Hickman, A. E., Poulton, A. J., Seeyave, S., and Lucas, M. I.: Iron-light interactions during the CROZet natural iron bloom and EXport experiment (CROZEX): II – Taxonomic responses and elemental stoichiometry, Deep Sea Res., 54, 2066–2084, 2007. </reference>
		<reference numeration="41" content_type="text"> Moore, J. K., Doney, S. C., and Lindsay, K.: Upper ocean ecosystem dynamics and iron cycling in a global three-dimensional model, Global Biogeochem. Cycles, 18, GB4028, doi:10.1029/2004GB002220, 2004. </reference>
		<reference numeration="42" content_type="text"> Moriceau, B., Soetaert, K., Gallinari, M., and Ragueneau, O.: Importance of particles formation to reconstruct water column biogenic silica fluxes, Global Biogeochem. Cycles, 21, GB3012, doi:10.1029/2006GB002814, 2007. </reference>
		<reference numeration="43" content_type="text"> Muggli, D. L., Lecourt, M., and Harrison, P. J.: Effects of iron and nitrogen source on the sinking rate, physiology and metal composition of an oceanic diatom from the subarctic Pacific, Mar. Ecol. Prog. Ser., 132, 215–227, 1996. </reference>
		<reference numeration="44" content_type="text"> Nelson, D. M., Tréguer, P., Brzezinski, M. A., Leynaert, A., and Quéguiner, B.: Production and dissolution of biogenic silica in the ocean, Revised global estimates, comparison with regional data and relationship to biogenic sedimentation, Global Biogeochem. Cycles, 9, 359–372, 1995. </reference>
		<reference numeration="45" content_type="text"> Nelson, D. M., Anderson, R. F., Barber, R. T., Brzezinski, M. A., Buesseler, K. O., Chase, Z., Collier, R. W., Dickson, M. L., François, R., Hiscock, M. R., Honjo, S., Marra, J., Martin, W. R., Sambrotto, R. N., Sayles, F. L., and Sigmon, D. E.: Vertical budgets for organic carbon and biogenic silica in the Pacific sector of the Southern Ocean, 1996–1998., Deep Sea Res., 49, 1645–1674, 2002. </reference>
		<reference numeration="46" content_type="text"> Peers, G., Quesnel, S.-A., and Price, N. M.: Copper requirements for iron acquisition and growth of coastal and oceanic diatoms, Limnol. Oceanogr., 50, 1149–1158, 2005. </reference>
		<reference numeration="47" content_type="text"> Peers, G. and Price, N. M.: Copper-containing plastocyanin used for electron transport by an oceanic diatom, Nature, 441, 341–344, 2006. </reference>
		<reference numeration="48" content_type="text"> Petroutsos, D., Terauchi, A. M., Busch, A., Hirschmann, I., Merchant, S. S., Finazzi, G., and Hippler, M.: PGRL1 Participates in Iron-induced Remodeling of the Photosynthetic Apparatus and in Energy Metabolism in \textitChlamydomonas reinhardtii, J. Biol. Chem., 284, 32770–32781, 2009. </reference>
		<reference numeration="49" content_type="text"> Pondaven, P., Fravalo, C., Ruiz-Pino, D., Tréguer, P., Quéguiner, B., and Jeandel, C.: Modelling the silica pump in the Permanently Open Ocean Zone of the Southern Ocean, J. Marine Syst., 17, 587–619, 1998. </reference>
		<reference numeration="50" content_type="text"> Pondaven, P., Ragueneau, O., Tréguer, P., Hauvespre, A., Dezileau, L., and Reyss, J.-L.: Resolving the &apos;opal pradox&apos; in the Southern Ocean, Nature, 405, 168–172, 2000. </reference>
		<reference numeration="51" content_type="text"> Pondaven, P., Gallinari, M., Chollet, S., Bucciarelli, E., Sarthou, G., Schultes, S., and Jean, F.: Grazing-induced changes in cell wall silicification in a marine diatom, Protist, 158, 21–28, 2007. </reference>
		<reference numeration="52" 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, 443–461, 1988/1989. </reference>
		<reference numeration="53" content_type="text"> Price, N. M.: The elemental stoichiometry and composition of an iron-limited diatom, Limnol. Oceanogr., 50, 1159–1171, 2005. </reference>
		<reference numeration="54" content_type="text"> Ragueneau, O. and Tréguer, P.: Determination of biogenic silica in coastal waters : applicability and limits of the alkaline digestion method, Mar. Chem., 45, 43–51, 1994. </reference>
		<reference numeration="55" content_type="text"> Rijstenbil, J. W. and Gerringa, L. J. A.: Interactions of algal ligands, metal complexation and availability, and cell responses of the diatom \textitDitylum brightwellii with a gradual increase in copper, Aquat. Toxic., 56, 115–131, 2002. </reference>
		<reference numeration="56" content_type="text"> Rogerson, A., DeFreitas, S. W., and McInnes, A. G.: Growth rates and ultrastructure of siliceous setae of \textitChaetoceros gracilis (Baccilariophyceae), J. Phycol., 22, 56–62, 1986. </reference>
		<reference numeration="57" content_type="text"> Sarthou, G., Timmermans, K. R., Blain, S., and Tréguer, P.: Growth physiology and fate of diatoms in the ocean: a review, J. Sea Res., 53, 25–42, 2005. </reference>
		<reference numeration="58" content_type="text"> Sigmon, D. E., Nelson, D. M., and Brzezinski, M. A.: The Si cycle in the Pacific sector of the Southern Ocean: seasonal diatom production in the surface layer and export to the deep sea, Deep Sea Res., 49, 1747–1763, 2002. </reference>
		<reference numeration="59" content_type="text"> Smetacek, V.: Diatoms and the Ocean Carbon Cycle, Protist, 150, 25–32, 1999. </reference>
		<reference numeration="60" content_type="text"> Strzepek, R. F. and Price, N. M.: Influence of irradiance and temperature on the iron content of the marine diatom \textitThalassiosira weissflogii (Bacillariophyceae), Mar. Ecol. Prog. Ser., 206, 107–117, 2000. </reference>
		<reference numeration="61" content_type="text"> Strzepek, R. F. and Harrison, P. J.: Photosynthetic architecture differs in coastal and oceanic diatoms, Nature, 431, 689–692, 2004. </reference>
		<reference numeration="62" content_type="text"> Sunda, W. G., Swift, D. G., and Huntsman, S. A.: Low iron requirement for growth in oceanic phytoplankton, Nature, 351, 55–57, 1991. </reference>
		<reference numeration="63" content_type="text"> Sunda, W. G. and Huntsman, S. A.: Iron uptake and growth limitation in oceanic and coastal phytoplankton, Mar. Chem., 50, 189–206, 1995. </reference>
		<reference numeration="64" content_type="text"> Sunda, W. G. and Huntsman, S. A.: Interrelated influence of iron, light and cell size on marine phytoplankton growth, Nature, 390, 389–392, 1997. </reference>
		<reference numeration="65" content_type="text"> Takeda, S.: Influence of iron availability on nutrient consumption ratio of diatoms in oceanic waters, Nature, 393, 774–777, 1998. </reference>
		<reference numeration="66" content_type="text"> Thornton, D. C. O.: Diatom aggregation in the sea: mechanisms and ecological implications., Eur. J. Phycol., 37, 149–161, 2002. </reference>
		<reference numeration="67" content_type="text"> Timmermans, K. R., Stolte, W., and de Baar, H. J. W.: Iron-mediated effects on nitrate reductase in marine phytoplankton, Mar. Biol., 121, 389–396, 1994. </reference>
		<reference numeration="68" content_type="text"> Timmermans, K. R., Davey, M. S., van der Wagt, B., Snoek, J., Geider, R. J., Veldhuis, M. J. W., Gerringa, L. J. A., and de Baar, H. J. W.: Co-limitation by iron and light of \textitChaetoceros brevis, \textitC. dichaeta and \textitC. calcitrans (Bacillariophyceae), Mar. Ecol. Prog. Ser., 217, 287–297, 2001a. </reference>
		<reference numeration="69" 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., Croot, P. L., and Boye, M.: Growth rates of large and small Southern Ocean diatoms in relation to availability of iron in natural seawater, Limnol. Oceanogr., 46, 260–266, 2001b. </reference>
		<reference numeration="70" content_type="text"> Timmermans, K. R., van der Wagt, B., and de Baar, H. J. W.: Growth rates, half-saturation constants, and silicate, nitrate, and phosphate depletion in relation to iron availability of four large, open ocean diatoms from the Southern Ocean, Limnol. Oceanogr., 49, 2141–2151, 2004. </reference>
		<reference numeration="71" content_type="text"> Timmermans, K. R., Veldhuis, M. J. W., and Brussaard, C. P. D.: Cell death in three marine diatom species in response to different irradiance levels, silicate, or iron concentrations, Aquat. Microb. Ecol., 46, 253–261, 2007. </reference>
		<reference numeration="72" content_type="text"> Waite, A. M., Thompson, P. A., and Harrison, P. J.: Does energy control the sinking rates of marine diatoms?, Limnol. Oceanogr., 37, 468–477, 1992. </reference>
	</references>
</article>
