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<article language="en">
	<journal>
		<journal_title>Nonlinear Processes  in Geophysics</journal_title>
		<journal_url>www.nonlin-processes-geophys.net</journal_url>
		<issn>1023-5809</issn>
		<eissn>1607-7946</eissn>
		<volume_number>16</volume_number>
		<issue_number>5</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/npg-16-623-2009</doi>
	<article_url>http://www.nonlin-processes-geophys.net/16/623/2009/</article_url>
	<abstract_html>http://www.nonlin-processes-geophys.net/16/623/2009/npg-16-623-2009.html</abstract_html>
	<fulltext_pdf>http://www.nonlin-processes-geophys.net/16/623/2009/npg-16-623-2009.pdf</fulltext_pdf>
	<start_page>623</start_page>
	<end_page>630</end_page>
	<publication_date>2009-10-30</publication_date>
	<article_title content_type="html">Plankton bloom controlled by horizontal stirring</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>W. McKiver</name>
			<email>william.mckiver@ucd.ie</email>
		</author>
		<author numeration="2" affiliations="1">
			<name>Z. Neufeld</name>
		</author>
		<author numeration="3" affiliations="2">
			<name>I. Scheuring</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">School of Mathematical Sciences &amp; Complex and Adaptive Systems Laboratory, University College Dublin, Ireland</affiliation>
		<affiliation numeration="2" content_type="html">Institute of Biology, Department of Plant Taxonomy and Ecology, Research Group of Theoretical Biology and Ecology, Eötvös University, Budapest, Hungary</affiliation>
	</affiliations>
	<abstract content_type="html">Here we show a simple mechanism in which changes in the rate of
horizontal stirring by mesoscale ocean eddies can trigger or
suppress plankton blooms and can lead to an abrupt change in the
average plankton density. We consider a single species phytoplankton
model with logistic growth, grazing and a spatially non-uniform
carrying capacity.  The local dynamics have multiple steady states
for some values of the carrying capacity that can lead to localized
blooms as fluid moves across the regions with different properties.
We show that for this model even small changes in the ratio of
biological timescales relative to the flow timescales can greatly
enhance or reduce the global plankton productivity. Thus, this may
be a possible mechanism in which changes in horizontal mixing can
trigger plankton blooms or cause regime shifts in some oceanic
regions. Comparison between the spatially distributed model and
Lagrangian simulations considering temporal fluctuations along fluid
trajectories, demonstrates that small scale transport processes also
play an important role in the development of plankton blooms with a
significant influence on global biomass.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Abraham, E. R.: The generation of plankton patchiness by turbulent stirring, Nature, 391, 577–580, 1998. </reference>
		<reference numeration="2" content_type="text"> Abraham, E. R., Law, C. S., Boyd, P. W., Lavender, S. J., Maldonado, M. T., and Bowie, A. R.: Importance of stirring in the development of an iron-fertilized phytoplankton bloom. Nature, 407, 727–730, 2000. </reference>
		<reference numeration="3" content_type="text"> Amemiya, T., Enomoto, T., Rossberg, A. G., Yamamoto, T., Inamori, Y., and Kiminori, I.: Stability and dynamical behavior in a lake-model and implications for regime shifts in real lakes Ecological Modelling, 206, 54–62, 2008. </reference>
		<reference numeration="4" content_type="text"> Bartello, P. and Thomas, S.: The cost-effectiveness of semi-Lagrangian advection, Mon. Weather Rev., 124, 2883–2897, 1996. </reference>
		<reference numeration="5" content_type="text"> Birch, D. A., Tsang, Y. K., and Young, W. R.: Bounding biomass in the Fisher equation, Phys. Rev. E, 75, 066304, doi:10.1103/PhysRevE.75.066304, 2007. </reference>
		<reference numeration="6" content_type="text"> Bracco, A., Provenzale, A., and Scheuring, I.: Mesoscale vortices and the paradox of the plankton, Proc. R. Soc. Lond. B, 267, 1795–1800, 2000. </reference>
		<reference numeration="7" content_type="text"> Collie, J. S., Richardson, K., and Steele, J. H.: Regime shifts: can ecological theory illuminate the mechanisms?, Prog. Oceanogr., 60, 281–302, 2004. </reference>
		<reference numeration="8" content_type="text"> deYoung, B., Barange, M., Beaugrand, G., Harris, R., Perry, R. I., Scheffer, M., and Werner, F.: Regime shifts in marine ecosystems: detection, prediction and management, Trends Ecol. Evol., 23, 402–409, 2008. </reference>
		<reference numeration="9" content_type="text"> Dritschel, D., Polvani, L., and Mohebalhojeh, A.: The contour-advective semi-Lagrangian algorithm for the shallow water equations, Mon. Weather Rev., 127, 1551–1565, 1999. </reference>
		<reference numeration="10" content_type="text"> Edwards, A. M. and Yool, A.: The role of higher predation in plankton population models, J. Plank. Res., 22, 1085–1112, 2000. \bibitem[Franke et al.(1999)] Franke-etal99 Franke, U., Hutter, K., and John, K.: A physical-biological coupled model for algal dynamics in lakes, B. Math. Biol., 61, 239–272, 1999. </reference>
		<reference numeration="11" content_type="text"> Hernández-Garc\&apos;ia, E., López, C., and Neufeld, Z.: Small-scale structure of nonlinearly interacting species advected by chaotic flows, Chaos, 12(2), 470–480, 2002. </reference>
		<reference numeration="12" content_type="text"> Huisman, J., Arrayás M., Ebert, U., and Sommeijer, B.: How do sinking phytoplankton species manage to persist?, American Naturalist, 159, 245–254, 2002. \bibitem[Huisman et al.(2006)] Huisman-etal06 Huisman, J., Nga, N., Thi, P., Karl, D. M., and Sommeijer, B.: Reduced mixing generates oscillations and chaos in the oceanic deep chlorophil maximum, Nature, 439, 322–325, 2006. \bibitem[Huppert et al.(2002)] Huppert-etal02 Huppert, A., Blasius, B., and Stone, L.: A model of phytoplankton blooms. American Naturalist, 159, 156–171, 2002. \bibitem[Huppert et al.(2005)] Huppert-etal05 Huppert, A., Blasius, B., Olinky, R., and Stone, L.: A model for seasonal phytoplankton blooms, J. Theor. Biol., 236, 276–290, 2005. </reference>
		<reference numeration="13" content_type="text"> Károlyi, G., Péntek, \&apos;A., Scheuring, I., Tél, T., and Toroczkai, Z.: Chaotic flow: the physics of species coexistence, Proc. Natl. Acad. Sci. USA, 97(25), 13661–13665, 2000. </reference>
		<reference numeration="14" content_type="text"> Lévy, M., Klein, P., and Treguier, A.: Impact of sub-mesoscale physics on production and subduction of phytoplankton in an oligotrophic regime, J. Mar. Res., 59, 535–565, 2001. </reference>
		<reference numeration="15" content_type="text"> Maltrud, M. E. and Vallis, G. K.: Energy spectra and coherent structures in forced two-dimensional and beta-plane turbulence, J. Fluid Mech., 228, 321–342, 1991. </reference>
		<reference numeration="16" content_type="text"> Martin, A. P., Richards, K., Bracco, A., and Provenzale, A.: Patchy productivity in the open ocean. Global Biogeochem. Cycles, 16, 2, doi:10.1029/2001GB001449, 2002. </reference>
		<reference numeration="17" content_type="text"> Martin, A.: Phytoplankton patchiness: the role of lateral stirring and mixing, Prog. Oceanogr., 57, 125–174, 2003. </reference>
		<reference numeration="18" content_type="text"> McKiver, W. and Neufeld, Z.: The influence of turbulent advection on the statistical properties of plankton ecosystems, Phys. Rev. E, 79, 061902, doi:10.1103/PhysRevE.79.061902, 2009. </reference>
		<reference numeration="19" content_type="text"> Miller, C.: Biological Oceanography, Blackwell Publ., UK, 2004. </reference>
		<reference numeration="20" content_type="text"> Neufeld, Z., Haynes, P., Garçon, V., and Sudre, J.: Ocean fertilization experiments may initiate a large scale phytoplankton bloom, Geophys. Res. Lett., 29(11), 1534, doi:10.1029/2001GL013677, 2002. </reference>
		<reference numeration="21" content_type="text"> Pasquero, C.: Differential eddy diffusion of biogeochemical tracers, Geophys. Res. Lett., 32, L17603, doi:10.1029/2005GL023662, 2005. </reference>
		<reference numeration="22" content_type="text"> Pasquero, C., Bracco, A., and Provenzale, A.: Impact of spatio-temporal variability of the nutrient flux on primary productivity in the ocean, J. Geophys. Res., 110, C07005, doi:10.1029/2004JC002738, 2005. </reference>
		<reference numeration="23" content_type="text"> Rossi, V., López, C., Sudre, J., Hernández-Garc\&apos;ia, E., and Garçon, V.: Comparative study of mixing and biological activity of the Benguela and Canary upwelling systems, Geophys. Res. Lett., 35, L11602, doi:10.1029/2008GL033610, 2008. </reference>
		<reference numeration="24" content_type="text"> Sandulescu, M., López, C., Hernández-García, E., and Feudel, U.: Plankton blooms in vortices: the role of biological and hydrodynamic timescales, Nonlin. Processes Geophys., 14, 443–454, 2007. </reference>
		<reference numeration="25" content_type="text"> Scheffer, M.: Multiplicity of stable states in freshwater systems, Hydrobiologia, 200/201, 475–486, 1990. </reference>
		<reference numeration="26" content_type="text"> Scheffer, M., Hosper, S. H., Meijer, M. L., and Moss, B.: Alternative equilibria in shallow lakes, Trends. Ecol. Evol., 8, 275–279, 1993. </reference>
		<reference numeration="27" content_type="text"> Scheffer, M., Carpenter, S., Folley, J. A., Folke, C., and Walker, B.: Catastrophic shifts in ecosystems, Nature, 413, 591–596, 2001. </reference>
		<reference numeration="28" content_type="text"> Steele, J. H. and Henderson, E. W.: The role of predation in plankton models, J. Plank. Res., 14, 157–172, 1992. </reference>
		<reference numeration="29" content_type="text"> Stone, L. and Berman, T.: Positive feedback in aquatic ecosystems: the case of microbial loop, B. Math. Biol., 55, 919–936, 1993. </reference>
		<reference numeration="30" content_type="text"> Tabeling, P.: Two-dimensional turbulence: a physicist approach, Phys. Reports, 362, 1–62, 2002. </reference>
		<reference numeration="31" content_type="text"> Temperton, C. and Staniforth, A.: An efficient two-time-level semi-Lagrangian semi implicit integrating scheme, Q. J. Roy. Meteorol. Soc., 113, 1025–1040, 1987. </reference>
		<reference numeration="32" content_type="text"> Truscott, J. E.: Environmental forcing of simple plankton models, J. Plank. Res., 17, 2207–2232, 1995. </reference>
		<reference numeration="33" content_type="text"> Yentsch, C. S., Lapointe, B. E., Poulton, N., and Phinney, D. A.: Anatomy of a red tide bloom off the southwest coast of Florida, Harmful Algae, 7(6), 817–826, 2008. \bibitem[Yoshiyama and Nakajima(2002)] Yoshiyama02 Yoshiyama, K. and Nakajima, H.: Catastrophic Transition in Vertical Distributions of Phytoplankton: Alternative Equilibria in a Water Column, J. Theor. Biol., 216, 397–408, 2002. </reference>
	</references>
</article>

