<?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>4</volume_number>
		<issue_number>5</issue_number>
		<publication_year>2007</publication_year>
	</journal>
	<doi>10.5194/bg-4-853-2007</doi>
	<article_url>http://www.biogeosciences.net/4/853/2007/</article_url>
	<abstract_html>http://www.biogeosciences.net/4/853/2007/bg-4-853-2007.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/4/853/2007/bg-4-853-2007.pdf</fulltext_pdf>
	<start_page>853</start_page>
	<end_page>868</end_page>
	<publication_date>2007-10-16</publication_date>
	<article_title content_type="html">Relationship between photosynthetic parameters and different proxies of phytoplankton biomass in the subtropical ocean</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>Y. Huot</name>
			<email>huot@obs-vlfr.fr</email>
		</author>
		<author numeration="2" affiliations="1">
			<name>M. Babin</name>
		</author>
		<author numeration="3" affiliations="2">
			<name>F. Bruyant</name>
		</author>
		<author numeration="4" affiliations="4">
			<name>C. Grob</name>
		</author>
		<author numeration="5" affiliations="3">
			<name>M. S. Twardowski</name>
		</author>
		<author numeration="6" affiliations="1">
			<name>H. Claustre</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">CNRS, Laboratoire d&apos;Océanographie de Villefranche, 06230 Villefranche-sur-Mer, France; Université Pierre et Marie Curie-Paris 6, Laboratoire d&apos;Océanographie de Villefranche, 06230 Villefranche-sur-Mer, France</affiliation>
		<affiliation numeration="2" content_type="html">Dalhousie University, Department of Oceanography, 1355 Oxford Street, Halifax N.S. B3H 4J1, Canada</affiliation>
		<affiliation numeration="3" content_type="html">WET Labs, Inc., Department of Research, 165 Dean Knauss Dr., Narragansett, RI 02882, USA</affiliation>
		<affiliation numeration="4" content_type="html">Graduate Program in Oceanography, Department of Oceanography and Center for Oceanographic Research in the eastern South Pacific, University of Concepción, Casilla 160-C, Concepción, Chile</affiliation>
	</affiliations>
	<abstract content_type="html">Probably because it is a readily available ocean color product, almost all
models of primary productivity use chlorophyll as their index of
phytoplankton biomass. As other variables become more readily available,
both from remote sensing and in situ autonomous platforms, we should ask if
other indices of biomass might be preferable. Herein, we compare the
accuracy of different proxies of phytoplankton biomass for estimating the
maximum photosynthetic rate (&lt;I&gt;P&lt;/I&gt;&lt;sub&gt;max&lt;/sub&gt;) and the initial slope of the
production versus irradiance (P vs. E) curve (α). The proxies
compared are: the total chlorophyll &lt;I&gt;a&lt;/I&gt; concentration (Tchl&lt;I&gt;a&lt;/I&gt;, the sum of
chlorophyll &lt;I&gt;a&lt;/I&gt; and divinyl chlorophyll), the phytoplankton absorption
coefficient, the phytoplankton photosynthetic absorption coefficient, the
active fluorescence in situ, the particulate scattering coefficient at 650 nm
(&lt;I&gt;b&lt;sub&gt;p&lt;/sub&gt;&lt;/I&gt;(650)), and the particulate backscattering
coefficient at 650 nm (&lt;I&gt;b&lt;sub&gt;bp&lt;/sub&gt;&lt;/I&gt;(650)). All of the data
(about 170 P vs. E curves) were collected in the South Pacific Ocean. We
find that when only the phytoplanktonic biomass proxies are available, &lt;I&gt;b&lt;sub&gt;p&lt;/sub&gt;&lt;/I&gt;(650) 
and Tchl&lt;I&gt;a&lt;/I&gt; are respectively the best estimators of
&lt;I&gt;P&lt;/I&gt;&lt;sub&gt;max&lt;/sub&gt; and α. When additional variables are available, such as
the depth of sampling, the irradiance at depth, or the temperature, Tchl&lt;I&gt;a&lt;/I&gt; is
the best estimator of both &lt;I&gt;P&lt;/I&gt;&lt;sub&gt;max&lt;/sub&gt; and α.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Babin, M. and Morel, A.: An incubator designed for extensive and sensitive measurements of phytoplankton photosynthetic parameters, Limnol. Oceanogr., 39, 694&amp;ndash;702, 1994. </reference>
		<reference numeration="2" content_type="text"> Babin, M., Morel, A., Claustre, H., Bricaud, A., Kolber, Z., and Falkowski, P. G.: Nitrogen- and irradiance-dependent variations of the maximum quantum yield of carbon fixation in eutrophic, mesotrophic and oligotrophic marine systems, Deep-Sea Res. Pt I, 43, 1241&amp;ndash;1272, 1996. </reference>
		<reference numeration="3" content_type="text"> Behrenfeld, M. J. and Falkowski, P. G.: A consumer&apos;s guide to phytoplankton primary productivity models, Limnol. Oceanogr., 42, 1479&amp;ndash;1491, 1997. </reference>
		<reference numeration="4" content_type="text"> Behrenfeld, M. J. and Boss, E.: The beam attenuation to chlorophyll ratio: An optical index of phytoplankton physiology in the surface ocean?, Deep-Sea Res. Pt I, 50, 1537&amp;ndash;1549, 2003. </reference>
		<reference numeration="5" content_type="text"> Behrenfeld, M. J., Prasil, O., Babin, M., and Bruyant, F.: In search of a physiological basis for covariations in light-limited and light-saturated photosynthesis, J. Phycol., 40, 4&amp;ndash;25, 2004. </reference>
		<reference numeration="6" content_type="text"> Behrenfeld, M. J., Boss, E., Siegel, D. A., and Shea, D. M.: Carbon-based ocean productivity and phytoplankton physiology from space, Global Biogeochem. Cy., 19, GB1006, doi:1010.1029/2004GB002299, 2005. </reference>
		<reference numeration="7" content_type="text"> Behrenfeld, M. J. and Boss, E.: Beam attenuation and chlorophyll concentration as alternative optical indices of phytoplankton biomass, J. Mar. Res., 64, 431&amp;ndash;451, 2006. </reference>
		<reference numeration="8" content_type="text"> Berges, J. A.: Ratios, regression statistics, and &quot;Spurious&quot; Correlation, Limnol. Oceanogr., 42, 1006&amp;ndash;1007, 1997. </reference>
		<reference numeration="9" content_type="text"> Bricaud, A., Claustre, H., Ras, J., and Oubelkheir, K.: Natural variability of phytoplankton absorption in oceanic waters: Influence of the size structure of algal populations, J. Geophys. Res.-Oceans, 109, C11010, doi:11010.11029/12004JC002419, 2004. </reference>
		<reference numeration="10" content_type="text"> Claustre, H., Bricaud, A., Babin, M., Bruyant, F., Guillou, L., Le Gall, F., Marie, D., and Partensky, F.: Diel variations in prochlorococcus optical properties, Limnol. Oceanogr., 47, 1637&amp;ndash;1647, 2002. </reference>
		<reference numeration="11" content_type="text"> Claustre, H., Huot, Y., Obernosterer, I., Gentili, B., Tailliez, D., Lewis, M.: Gross community production and metabolic balance in the South Pacific Gyre, using a non intrusive bio-optical method, Biogeosciences Discuss., 4, 3089&amp;ndash;3121, 2007. </reference>
		<reference numeration="12" content_type="text"> Falkowski, P. G. and Raven, J. A.: Aquatic photosynthesis, First ed., Blackwell Science, Malden, 375 pp., 1997. </reference>
		<reference numeration="13" content_type="text"> Gardner, W. D., Mishonov, A., and Richardson, M. J.: Global poc concentrations from in-situ and satellite data, Deep-Sea Res. Part II, 53, 718&amp;ndash;740, 2006. </reference>
		<reference numeration="14" content_type="text"> Gregg, W. W. and Carder, K. L.: A simple spectral solar irradiance model for cloudless maritime atmosphere, Limnol. Oceanogr., 35, 1657&amp;ndash;1675, 1990. </reference>
		<reference numeration="15" content_type="text"> Grob, C., Ulloa, O., Claustre, H., Huot, Y., Alarcon, 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="16" content_type="text"> Kitchen, J. C., Ronald, J., and Zaneveld, V.: On the noncorrelation of the vertical structure of light scattering and chlorophyll a in case I waters, J. Geophys. Res.-Oceans, 95, 20 237&amp;ndash;20 246, 1990. </reference>
		<reference numeration="17" content_type="text"> Lee, Z. P., Carder, K. L., Marra, J., Steward, R. G., and Perry, M. J.: Estimating primary production at depth from remote sensing, Appl. Optics, 35, 463&amp;ndash;474, 1996. </reference>
		<reference numeration="18" content_type="text"> Loisel, H. and Morel, A.: Light scattering and chlorophyll concentration in case 1 waters: A reexamination, Limnol. Oceanogr., 43, 847&amp;ndash;858, 1998. </reference>
		<reference numeration="19" content_type="text"> Longhurst, A. R.: Ecological geography of the sea, Academic Press, San Diego, 398 pp., 1998. </reference>
		<reference numeration="20" 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: Anderson, R. M., Academic Press, 2005. </reference>
		<reference numeration="21" content_type="text"> Marra, J., Trees, C. C., and O&apos;Reilly, J. E.: Phytoplankton pigment absorption: A strong predictor of primary productivity in the surface ocean, Deep-Sea Res. Pt I, 54, 155&amp;ndash;163, 2007. </reference>
		<reference numeration="22" content_type="text"> Morel, A.: Diffusion de la lumière par les eaux de mer. Resultats expérimentaux et approche théorique., AGARD lectures series, 3.1.1&amp;ndash;3.1.76, 1973. </reference>
		<reference numeration="23" content_type="text"> Morel, A. and Bricaud, A.: Inherent properties of algal cells including picoplankton: Theoretical and experimental results, in: Photosynthetic picoplankton, edited by: Platt, T. and Li, W. K. W., Canadian bulletin of fisheries and aquatic sciences, 521&amp;ndash;559, 1986. </reference>
		<reference numeration="24" content_type="text"> Morel, A.: Optical modeling of the upper ocean in relation to its biogenous matter content (case 1 waters), J. Geophys. Res.-Oceans, 93, 10 749&amp;ndash;10 768, 1988. </reference>
		<reference numeration="25" content_type="text"> Morel, A. and Ahn, Y.-H.: Optics of heterotrophic nanoflagellates and ciliates: A tentative assessment of their scattering role in oceanic waters compared to those of bacterial and algal cells, J. Mar. Res., 49, 177&amp;ndash;202, 1991. </reference>
		<reference numeration="26" content_type="text"> Morel, A. and Maritorena, S.: Bio-optical properties of oceanic waters: A reappraisal, J. Geophys. Res.-Oceans, 106, 7163&amp;ndash;7180, 2001. </reference>
		<reference numeration="27" content_type="text"> Morel, A., Gentili, B., Claustre, H., Babin, M., Bricaud, A., Ras, J., and Tieche, F.: Optical properties of the &quot;Clearest&quot; Natural waters, Limnol. Oceanogr., 52, 217&amp;ndash;229, 2007. </reference>
		<reference numeration="28" content_type="text"> Oubelkheir, K., Claustre, H., Sciandra, A., and Babin, M.: Bio-optical and biogeochemical properties of different trophic regimes in oceanic waters, Limnol. Oceanogr., 50, 1795&amp;ndash;1809, 2005. </reference>
		<reference numeration="29" content_type="text"> Perry, M. J.: Measurements of phytoplankton absorption other than per unit of chlorophyll a, in: Ocean optics, edited by: Spinrad, R. W., Carder, K. L., and Perry, M. J., Oxford monographs on geology and geophysics, Oxford University Press, New York, 107&amp;ndash;116, 1994. </reference>
		<reference numeration="30" content_type="text"> Platt, T., Gallegos, C. L., and Harrison, W. G.: Photoinhibition of photosynthesis in natural assemblages of marine phytoplankton, J. Mar. Res., 38, 687&amp;ndash;701, 1980. </reference>
		<reference numeration="31" content_type="text"> Platt, T. and Sathyendranath, S.: Spatial structure of pelagic ecosystem processes in the global ocean, Ecosystems, 2, 384&amp;ndash;394, 1999. </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 Discuss., 4, 3409&amp;ndash;3451, 2007. </reference>
		<reference numeration="33" content_type="text"> Siegel, D. A., Dickey, T. D., Washburn, L., Hamilton, M. K., and Mitchell, B. G.: Optical determination of particulate abundance and production variations in the oligtrophic ocean, Deep-Sea Res., 36, 211&amp;ndash;222, 1989. </reference>
		<reference numeration="34" content_type="text"> Sokal, R. R. and Rohlf, F. J.: Biometry the principles and practice of statistics in biological research, 3 ed., W.H. Freeman and Company, New York, 887 pp., 1995. </reference>
		<reference numeration="35" content_type="text"> Stramski, D. and Reynolds, R. A.: Diel variations in the optical properties of a marine diatom, Limnol. Oceanogr., 38, 1347&amp;ndash;1364, 1993. </reference>
		<reference numeration="36" content_type="text"> Stramski, D., Shalapyonok, A., and Reynolds, C. S.: Optical characterization of the marine oceanic unicellular cyanobacterium synechococcus grown under a day-night cycle in natural irradiance, J. Geophys. Res.-Oceans, 100, 13 295&amp;ndash;13 307, 1995. </reference>
		<reference numeration="37" content_type="text"> Stramski, D., Reynolds, C. S., Kahru, M., and Mitchell, B. G.: Estimation of particulate organic carbon in the ocean from remote sensing, Science, 285, 239&amp;ndash;242, 1999. </reference>
		<reference numeration="38" content_type="text"> Stramski, D., Boss, E., Bogucki, D., and Voss, K. J.: The role of seawater constituents in light backscattering in the ocean, Prog. Oceanogr., 61, 27&amp;ndash;56, 2004. </reference>
		<reference numeration="39" content_type="text"> Sukenic, A., Bennett, J., and Falkowski, P. G.: Light-saturated photosynthesis &amp;ndash; limitation by electron transport or carbon fixation, Biochim. Biophys. Acta, 891, 205&amp;ndash;215, 1987. </reference>
		<reference numeration="40" content_type="text"> Sullivan, J. M., Twardowski, M. S., Zaneveld, J. R., Moore, C., Barnard, A., Donaghay, P. L., and Rhoades, B.: The hyperspectral temperature and salinity dependencies of absorption by water and heavy water in the 400&amp;ndash;750 nm spectral range, Appl. Optics, 45, 5294&amp;ndash;5309, 2006. </reference>
		<reference numeration="41" content_type="text"> Twardowski, M. S., Sullivan, J. M., Donaghay, P. L., and Zaneveld, J. R.: Microscale quantification of the absorption by dissolved and particulate material in coastal waters with an ac-9, J. Atmos. Ocean. Tech., 16, 691&amp;ndash;707, 1999. </reference>
		<reference numeration="42" content_type="text"> Twardowski, M. S., Claustre H., Freeman, A., Stramski, D., and Huot, Y.: Optical backscattering properties of the &quot;clearest&quot; natural waters, Biogeosciences Discuss., 4, 2441&amp;ndash;2491, 2007. </reference>
		<reference numeration="43" content_type="text"> Zaneveld, J. R. V., Kitchen, J. C., and Moore, C. C.: The scattering error correction of reflecting tube absorption meters, Ocean Optics XII, 44&amp;ndash;55, 1994. </reference>
	</references>
</article>

