Biogeosciences
www.biogeosciences.net
1726-4170
1726-4189
5
3
2008
10.5194/bg-5-707-2008
http://www.biogeosciences.net/5/707/2008/
http://www.biogeosciences.net/5/707/2008/bg-5-707-2008.html
http://www.biogeosciences.net/5/707/2008/bg-5-707-2008.pdf
707
718
2008-05-06
Build-up and decline of organic matter during PeECE III
K. G. Schulz
U. Riebesell
R. G. J. Bellerby
H. Biswas
M. Meyerhöfer
M. N. Müller
J. K. Egge
J. C. Nejstgaard
C. Neill
J. Wohlers
E. Zöllner
Leibniz Institute for Marine Sciences (IFM-GEOMAR), Düsternbrooker Weg 20, 24105 Kiel, Germany
Bjerknes Centre for Climate Research, University of Bergen, Allégaten 55, 5007 Bergen, Norway
Geophysical Institute, University of Bergen, Allégaten 70, 5007 Bergen, Norway
University of Bergen, Department of Biology, Box 7800, 5020 Bergen, Norway
Increasing atmospheric carbon dioxide (CO<sub>2</sub>) concentrations due to
anthropogenic fossil fuel combustion are currently changing the ocean's
chemistry. Increasing oceanic [CO<sub>2</sub>] and consequently
decreasing seawater pH have the potential to significantly impact
marine life. Here we describe and analyze the build-up and decline of a
natural phytoplankton bloom initiated during the 2005 mesocosm
Pelagic Ecosystem CO<sub>2</sub> Enrichment study (PeECE III). The draw-down
of inorganic nutrients in the upper surface layer of the mesocosms was
reflected by a concomitant increase of organic matter until day
<i>t</i><sub>11</sub>, the peak of the bloom. From then on, biomass standing stocks
steadily decreased as more and more particulate organic matter was
lost into the deeper layer of the mesocosms. We show that organic
carbon export to the deeper layer was significantly enhanced at
elevated CO<sub>2</sub>. This phenomenon might have impacted organic matter
remineralization leading to decreased oxygen concentrations in the
deeper layer of the high CO<sub>2</sub> mesocosms as indicated by
deep water ammonium concentrations. This would have
important implications for our understanding of pelagic ecosystem
functioning and future carbon cycling.
Allgaier, M., Riebesell, U., Vogt, M., Thyrhaug, R., and Grossart, H.-P.: Coupling of heterotrophic bacteria to phytoplankton bloom development at different $p$CO<sub>2</sub> levels: a mesocosm study, Biogeosciences Discuss., 5, 317–359, 2008.
Antia, A. N., Suffrian, K., Holste, L., Müller, M. N., Nejstgaard, J. C., Simonelli, P., Carotenuto, Y., and Putzeys, S.: Dissolution of coccolithophorid calcite by microzooplankton and copepod grazing, Biogeosciences Discuss., 5, 1–23, 2008.
Arrigo K R.:Carbon cycle – Marine manipulations, Nature, 450, 491–492, 2007.
Barcelos~e Ramos, J., Biswas, H., Schulz, K G., LaRoche, J., and Riebesell, U.: Effect of rising atmospheric carbon dioxide on the marine nitrogen fixer \textitTrichodesmium, Global Biogeochem. Cy., 21, GB2028, doi:10.1029/2006GB002898, 2007.
Bellerby, R. G. J., Schulz, K. G., Riebesell, U., Neill, C., Nondal, G., Johannessen, T., and Brown, K. R.: Marine ecosystem community carbon and nutrient uptake stoichiometry under varying ocean acidification during the PeECE III experiment, Biogeosciences Discuss., 4, 4631–4652, 2007. %
%Breitbarth, E., Bellerby, R J., Neill, C., Ardelan, M V., Meyerhöfer, M., % Zöllner, E., Croot, P L., and Riebesell, U.: Ocean acidification % increases bioavailability of iron, Science (in review), 2007.\blackbox\bf % If still in review this has to be a footnote
Burkhardt, S. and Riebesell, U.: CO<sub>2</sub> availability affects elemental composition (C:N:P) of the marine diatom \textitSceletonema costatum, Mar. Ecol. Prog. Ser., 155, 67–76, 1997.
Burkhardt, S., Riebesell, U., and Zondervan, I.: Effects of growth rate, CO<sub>2</sub> concentration, and cell size on the stable carbon isotope fractionation in marine phytoplankton, Geochim. Cosmochim. Acta, 63, 3729–3741, 1999a.
Burkhardt, S., Zondervan, I., and Riebesell, U.: Effect of CO<sub>2</sub> concentration on the C:N:P ratio in marine phytoplankton: A species comparision, Limnol. Oceanogr., 44, 683–690, 1999b.
Caldeira, K. and Wickett, M E.: Anthropogenic carbon and ocean pH, Nature, 425, p 365, 2003.
Carotenuto, Y., Putzeys, S., Simonelli, P., Paulino, A., Meyerhöfer, M., Suffrian, K., Antia, A., and Nejstgaard, J. C.: Copepod feeding and reproduction in relation to phytoplankton development during the PeECE III mesocosm experiment, Biogeosciences Discuss., 4, 3913–3936, 2007.
Chapelle, A.: A preliminary model of nutrient cycling in sediments of a Mediterranean lagoon, Ecol. Mod., 80, 131–147, 1995.
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 pCO<sub>2</sub> during experimental blooms of the coccolithophorid \textitEmiliania huxleyi, Global Biogeochem. Cy., 19, GB2023, doi:10.1029/2004GB002318, 2005.
Egge, J K., Thingstad, T F., Engel, A., and Riebesell, U.: Primary production during nutrient-induced blooms at elevated CO<sub>2</sub> concentrations, Biogeosciences Discuss., 4, 4385–4410, 2007.
Engel, A.: Direct relationship between CO<sub>2</sub> uptake and transparent exopolymer particles production in natural phytoplankton, J. Plankt. Res., 24, 49–53, 2002.
Engel, A., Thoms, S., Riebesell, U., Rochelle-Newall, E., and Zondervan, I.: Polysaccharide aggregation as a potential sink of marine dissolved organic carbon, Nature, 428, 929–932, 2004.
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<sub>2</sub> concentration on a bloom of the coccolithophorid \textitEmiliania huxleyi in mesocosm experiments, Limnol. Oceanogr., 50, 493–507, 2005.
Gattuso, J.-P., Frankignoulle, M., Bourge, I., Romaine, S., and Buddemeier, R W.: Effect of calcium carbonate saturation of seawater on coral calcification, Global Planet. Change, 18, 37–46, 1998.
Hansen, H P. and Koroleff, F.: Determination of nutrients, in: Methods of seawater analysis, 3 Ed., edited by: Grasshoff, K., Kremling, K., and Ehrhardt, M., Wiley VCH, Weinheim, 159–228, 1999.
Holmes, R M., Aminot, A., Kérouel, R., Hooker, B A., and Peterson, B J.: A simple and precise method for measuring ammonium in marine and freshwater ecosystems, Can. J. Fish. Aquat. Sci., 56, 1801–1808, 1999.
Kleypas, J A., Buddemeier, R W., Archer, D., Gattuso, J.-P., Langdon, C., and Opdyke, B N.: Geochemical consequences of increased atmospheric carbon dioxide on coral reefs, Science, 284, 118–120, 1999.
Langer, G., Geisen, M., Baumann, K H., Kläs, J., Riebesell, U., Thoms, S., and Young, J R.: Species-specific responses of calcifying algae to changing seawater carbonate chemistry, Geochem. Geophys. Geosystems, 7, Q09006, doi:10.1029/2005GC001227, 2006.
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<sub>2</sub> levels, Biogeosciences, 5, 523–533, 2008.
Mackey, M D., Mackey, D J., Higgins, H W., and Wright, S W.: CHEMTAX – a program for estimating class abundances from chemical markers: application to HPLC measurements of phytoplankton, Mar. Ecol. Prog. Ser., 144, 265–283, 1996.
Meehl, G A., Stocker, T F., Collins, W D., Friedlingstein, P., Gaye, A T., Gregory, J M., Kitoh, A., Knutti, R., Murphy, J M., Noda, A., Raper, S. C B., Watterson, I G., Weaver, A J., and Zhao, Z.-C.: Global Climate Projections, 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 Kindom and new York, NY, USA, 747–845, 2007.
Orr, J. C Fabry, V J., Aumont, O., Bopp, L., Doney, S C., Feely, R A., Gnanadesikan, A., Gruber, N., Ishida, A., Joos, F., Key, R M., Lindsay, K., Maier-Reimer, E., Matear, R., Monfray, P., Mouchet, A., Najjar, R G., Plattner, G K., Rodgers, K B., Sabine, C L., Sarmiento, J L., Schlitzer, R., Slater, R D., Totterdell, I J., Weirig, M F., Yamanaka, Y., and Yool, A.: Anthropogenic ocean acidification over the twenty-first century and its impact on calcifying organisms, Nature, 437, 681–686, 2005.
Paulino, A. I, Egge, J. K., and Larsen, A.: Effects of increased atmospheric CO<sub>2</sub> on small and intermediate sized osmotrophs during a nutrient induced phytoplankton bloom, Biogeosciences Discuss., 4, 4173–4195, 2007.
Prentice, I C., Farquhar, G D., Fasham, M. J R., Goulden, M L., Heimann, M., Jaramillo, V J., Kheshgi, H S., Le~Quéré, C., Scholes, R J., and Wallace, D. W R.: The Carbon Cycle and Atmospheric Carbon Dioxide, in: Climate Change 2001: The Scientific Basis, Contribution of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change, edited by: Houghton, J. T., Ding, Y., Griggs, D. J., Nougher, M., van der Linden, P. J., Dai, X., Maskell, K., and Johnson, C. A., Cambridge University Press, Cambridge, United Kindom and new York, NY, USA, 183–237, 2001.
Qian, J. and Mopper, K.: Automated High-Performance, High-Temperature Combustion Total Organic Carbon Analyzer, Anal. Chem., 68, 3090–3097, 1996.
Riebesell, U.: Effects of CO<sub>2</sub> Enrichment on Marine Phytoplankton, J. Oceanogr., 60, 719–729, 2005. %
%Riebesell, U., Bellerby, R. G. J., Grossart, H.-P., and Thingstad, T. F.: %Mesocosm CO<sub>2</sub> pertubation studies: from organism to community level, Biogeosciences Discuss., in preparation, 2007.
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 atmosphe ric CO<sub>2</sub>, Nature, 407, 364–367, 2000.
Riebesell, U., Schulz, K G., Bellerby, R. G J., Botros, M., Fritsche, P., Meyerhöfer, N., Neill, C., G., N., Oschlies, A., Wohlers, J., and Zöllner, E.: Enhanced biological carbon consumption in a high CO<sub>2</sub> ocean, Nature, 450, 545–548, doi:10.1038/nature06267, 2007.
Riebesell, U., Bellerby, R. G. J., Grossart, H.-P., and Thingstad, F.: Mesocosm CO<sub>2</sub> perturbation studies: from organism to community level, Biogeosciences Discuss., 5, 641–659, 2008.
Rost, B., Zondervan, I., and Riebesell, U.: Light-dependent carbon isotope fractionation in the coccolithophorid \textitEmiliania huxleyi, Limnol. Oceanogr., 47, 120–128, 2002.
Russel, A D., Hönisch, B., Spero, H J., and Lea, D W.: Effects of seawater carbonate ion concentration and temperature on shell U, Mg, and Sr in cultured planktonic foraminifera, Geochim. Cosmochim. Acta, 68, 4347–4361, 2004.
Sarathchandra, S U.: Nitrification activities and the changes in the populations of nitrifying bacteria in soil perfused at two different H-ion concentrations, Plant Soil, 50, 99–111, 1978.
Serpa, D., Falcão, M., Duarte, P., Cancela~da Fonseca, L., and Vale, C.: Evaluation of ammonium and phosphate release from intertidal and subtidal sediments of a shallow coastal lagoon (Ria Formosa – Portugal): a modelling approach, Biogeochem., 82, 291–304, 2007.
Sinha, V., Williams, J., Meyerhöfer, M., Riebesell, U., Paulino, A I., and Larsen, A.: Air-sea fluxes of methanol, acetone, acetaldehyde, isoprene and DMS from a Norwegian fjord following a phytoplankton bloom in a mesocosm experiment, Atmos. Chem. Phys., 7, 739–755, 2007.
Strauss, E A., and Lamberti, G A.: Regulation of nitrification in aquatic sediments by organic carbon, Limnol. Oceanogr., 45, 1854–1859, 2000.
Suffrian, K., Simonelli, P., Nejstgaard, J. C., Putzeys, S., Carotenuto, Y., and Antia, A. N.: Microzooplankton grazing and phytoplankton growth in marine mesocosms with increased CO<sub>2</sub> levels, Biogeosciences Discuss., 5, 411–433, 2008.
Tanaka, T., Thingstad, T. F., Løvdal, T., Grossart, H.-P., Larsen, A., Schulz, K. G., and Riebesell, U.: Availability of phosphate for phytoplankton and bacteria and of labile organic carbon for bacteria at different $p$CO<sub>2</sub> levels in a mesocosm study, Biogeosciences Discuss., 4, 3937–3960, 2007.
Vogt, M., Steinke, M., Turner, S., Paulino, A., Meyerhöfer, M., Riebesell, U., Le~Quére, C., and Liss, P.: Dynamics of dimethylsulphoniopropionate and dimethylsulfide under different CO<sub>2</sub> concentrations during a mesocosm experiment, Biogeosciences, 4, 3673–3699, 2007a.
Vogt, M., Turner, S., Yassaa, N., Steinke, M., Williams, J., and Liss, P.: Laboratory inter-comparison of dissolved dimethyl sulphide (DMS) measurements using purge-and-trap and solid-phase microextraction techniques during a mesocosm experiment, Mar. Chem., 108, 32–39, 2008.
Wingenter, O W., Haase, K B., Zeigler, M., Blake, D R., Sherwood~Rowland, F S., Sive, B C., Paulino, A., Thyrhaug, R., Larsen, A., Schulz, K., Meyerhöfer, M., and Riebesell, U.: Unexpected consequences of increasing CO<sub>2</sub> and ocean acidity on marine production of DMS and CH<sub>2</sub>ClI: Potential climate impacts, Geophys. Res. Lett., 34, L05710, doi:10.1029/2006GL028139, 2007.