La turbulencia como mecanismo forzante de la distribución vertical del plancton en el océano subsuperficial
DOI:
https://doi.org/10.3989/scimar.03854.03APalabras clave:
turbulencia, máximo profundo de clorofila, distribución vertical del planctonResumen
Las distribuciones verticales de la tasa de disipación de la energía turbulenta y de la fluorescencia han sido medidas simultáneamente usando un microperfilador de alta resolución en cuatro regiones oceánicas distintas, desde zonas templadas a polares y desde regiones costeras al océano abierto. Valores altos de fluorescencia, conformando un máximo profundo de clorofila (MPC) fueron detectados habitualmente en regiones poco estratificadas de la columna de agua y justo por debajo de capas con valores máximos de la tasa de disipación de la energía turbulenta. En las cercanías del MPC se encontró una relación estadísticamente significativa y negativa entre los valores de fluorescencia y energía turbulenta. En este trabajo se discuten los posibles mecanismos que pueden explicar estas relaciones, incluyendo el efecto de la variación vertical del coeficiente de difusión y la posible alteración de la velocidad de sedimentación de las células de fitoplancton por el movimiento turbulento. Estos resultados proporcionan un mayor conocimiento sobre los procesos que controlan la distribución vertical de la comunidad pelágica y la posición de los MPC.
Descargas
Citas
Acuña J.L., López-Alvarez M., Nogeira E., González-Taboada F. 2010. Diatom flotation at the onset of the spring phytoplankton bloom: an in situ experiment. Mar. Ecol. Prog. Ser. 400: 115-125. http://dx.doi.org/10.3354/meps08405
Abbot M.R., Denman K.L., Powell T.M., Richerson P.J., Richards R.C., Goldman C.R. 1984. Mixing and the dynamics of the deep chlorophyll maximum in Lake Tahoe. Limnol. Oceanogr. 29: 862-878. http://dx.doi.org/10.4319/lo.1984.29.4.0862
Abraham E.R. 1998. The generation of plankton patchiness by turbulent stirring. Nature 391: 577-580. http://dx.doi.org/10.1038/35361
Aliseda A., Cartellier A., Hainaux F., Lasheras J.C. 2002. Effect of preferential concentration on the settling velocity of heavy particles in homogeneous isotropic turbulence. J. Fluid Mech. 468: 77-105. http://dx.doi.org/10.1017/S0022112002001593
Belyaev V.I. 1992. Modelling the influence of turbulence on phytoplankton photosynthesis. Ecol. Model. 60: 11-29. http://dx.doi.org/10.1016/0304-3800(92)90010-C
Bienfang P.K. 1980. Phytoplankton Sinking Rates in Oligotrophic Waters off Hawaii, USA. Mar. Biol. 61: 69-77. http://dx.doi.org/10.1007/BF00410342
Bosse T., Kleiser L., Miburg E. 2006. Small particles in homogeneous turbulence: Settling velocity enhancement by two-way coupling. Phys. Fluids 18: 027102. http://dx.doi.org/10.1063/1.2166456
Bowen J. D., Stolzenbach K. D., Chisholm S.W. 1993. Simulating bacterial clustering around phytoplankton cells in a turbulent ocean. Limnol. Oceanogr. 38: 36-51. http://dx.doi.org/10.4319/lo.1993.38.1.0036
Brainerd K.E., Gregg M.C. 1995. Surface mixed and mixing layer depths. Deep-Sea Res. I 42(9): 1521-1543. http://dx.doi.org/10.1016/0967-0637(95)00068-H
Brucato A., Grisafi F., Montante G. 1998. Particle drag coefficients in turbulent fluids. Chem. Eng. Sci. 53: 3295-3314. http://dx.doi.org/10.1016/S0009-2509(98)00114-6
Bruno M., Alonso J.J., Cozar A., Vidal J., Ruiz-Canavate A., Echevarria F., Ruiz J. 2002. The boiling water phenomena at Camarinal Sill, the Strait of Gibraltar. Deep-Sea Res. II 49: 4097-4113. http://dx.doi.org/10.1016/S0967-0645(02)00144-3
Carniel S., Kantha L.H., Book J.W., Sclavo M., Prandke H. 2012. Turbulence variability in the upper layers of the Southern Adriatic Sea under a variety of atmospheric forcings conditions. Cont. Shelf Res. 44: 39-56. http://dx.doi.org/10.1016/j.csr.2011.01.003
Cowles T.J., Desiderio R.A., Carr M.-E. 1998. Small-scale planktonic structure: Persistence and trophic consequences. Oceanogr. 11: 4-9. http://dx.doi.org/10.5670/oceanog.1998.08
Cullen J.J., Eppley R.W. 1981. Chlorophyll maximum layers of the Southern California Bight and possible mechanisms of their formation and maintenance. Oceanol. Acta 4: 23-32.
Cullen J.J. 1982. The deep chlorophyll maximum: comparing vertical profiles of chlorophyll a. Can. J. Fish. Aqua. Sci. 39: 791-803. http://dx.doi.org/10.1139/f82-108
Cullen J.J., Lewis M.R. 1988. The kinetics of algal photoadaptation in the context of vertical mixing. J. Plank. Res. 10(5): 1039-1063. http://dx.doi.org/10.1093/plankt/10.5.1039
D'Ovidio F., De Monte S., Alvain S., Dondonneau Y., Lévy M. 2010. Fluid dynamical niches of phytoplankton types. Proc. Natl. Acad. Sci. USA 107: 18366-18370. http://dx.doi.org/10.1073/pnas.1004620107
De Lillo F., Cecconi F., Lacorata G., Vulpiani A. 2008. Sedimentation speed of inertial particles in laminar and turbulent flows. EPL 84: 40005. http://dx.doi.org/10.1209/0295-5075/84/40005
Deksheniecks M.M., Donaghay P.L., Sullivan J.M., Rines J.E.B., Osborn T.R., Twardowski M.S. 2001. Temporal and spatial occurrence of thin phytoplankton layers in relation to physical processes. Mar. Ecol. Prog. Ser. 223: 61-71. http://dx.doi.org/10.3354/meps223061
Doubell M.J., Yamazaki H., Li H., Kokuby Y. 2009. An advanced laser-based fluorescence microstructure profiler (TurboMAP-L) for measuring bio-physical coupling in aquatic systems. J. Plank. Res. 31: 1441-1452. http://dx.doi.org/10.1093/plankt/fbp092
Druet C., Zielinski A. 1994. Modelling the fine-structure of the phytoplankton concentration in a stable stratified sea. Oceanol. Acta 17: 79-88.
Dzierzbicka L. 2006. Effects of turbulent mixing on the marine plankton vertical distribution: model simulations. Polish J. Ecol. 54: 215-230.
Estrada M., Marrase C., Latasa M., Berdalet E., Delgado M., Riera T. 1993. Variability of the Deep Chlorophyll Maximum characteristics in the northwestern Mediterranean. Mar. Ecol. Prog. Ser. 92: 289-300. http://dx.doi.org/10.3354/meps092289
Falkowski P.G. 1998. Light-shade adaptation in marine phytoplankton. In: Falkowski P.G. (ed.), Primary productivity in the sea. Plenum Press, pp. 99-119.
Falkowski P.G., Barber R., Smetacek V. 1998. Biogeochemical controls and feedbacks on ocean primary production. Science 281: 200-206. http://dx.doi.org/10.1126/science.281.5374.200
Falkowski P.G., Oliver M.J. 2007. Mix and match: how climate selects phytoplankton. Nature Rev. Microb. 5: 813-819. http://dx.doi.org/10.1038/nrmicro1751
Fasham M.J.R., Platt T., Irwin B., Jones K. 1985. Factors Affecting the Spatial Pattern of the Deep Chlorophyll Maximum in the Region of the Azores Front. Prog. Oceanogr. 14: 129-165. http://dx.doi.org/10.1016/0079-6611(85)90009-6
Fennel K., Boss E. 2003. Subsurface maxima of phytoplankton and chlorophyll, Steady-state solutions from a simple model. Limnol. Oceanogr. 48: 1521-1534. http://dx.doi.org/10.4319/lo.2003.48.4.1521
Franks P.J.S. 2005. Plankton patchiness, turbulent transport and spatial spectra. Mar. Ecol. Prog. Ser. 294: 295-309. http://dx.doi.org/10.3354/meps294295
Gould R.W. 1988. Net phytoplankton in a Gulf Stream warm-core ring: species composition, relative abundance, and the chlorophyll maximum layer. Deep-Sea Res. 35: 1595-1614. http://dx.doi.org/10.1016/0198-0149(88)90105-7
Gregg M.C., Alford M.H., Kontoyiannis H., Zervakis V., Winkel D. 2012. Mixing over the steep side of the Cycladic Plateau in the Aegean Sea. J. Mar. Syst. 89: 30-47. http://dx.doi.org/10.1016/j.jmarsys.2011.07.009
Herbland A., Voituriez B. 1979. Hydrological structure analysis for estimating the primary production in the tropical Atlantic Ocean. J. Mar. Res. 37: 87-101.
Hodges B.A., Rudnick D.L. 2004. Simple models of steady deep maxima in chlorophyll and biomass. Deep-Sea Res. I 51: 999-1015. http://dx.doi.org/10.1016/j.dsr.2004.02.009
Husiman J., Pham-Ti N.N., Karl D.M., Sommeijer B. 2006. Reduced mixing generates oscillations and chaos in the oceanic deep chlorophyll maximum. Nature 439(7074): 322-325. http://dx.doi.org/10.1038/nature04245
Jassby A., Powell T.M. 1975. Vertical patterns of eddy diffusion during stratification in Castle Lake, California. Limnol. Oceanogr. 20: 530-543. http://dx.doi.org/10.4319/lo.1975.20.4.0530
Jumars P.A., Trowbrige J.H., Boss E., Karp-Boss L. 2009. Turbulence-plankton interactions: a new cartoon. Mar. Ecol. 30: 133-150. http://dx.doi.org/10.1111/j.1439-0485.2009.00288.x
Karl D.M., Holm-Hasen O., Taylor G.T., Tien G., Bird D.F. 1991. Microbial biomass and productivity in the western Bransfield Strait, Antarctica during the 1986-87 austral summer. Deep-Sea Res. II 38: 1029-1055.
Kiørboe, T. 1993. Turbulence, phytoplankton cell size, and the structure of pelagic food webs. Adv. Mar. Biol. 29: 1-72. http://dx.doi.org/10.1016/S0065-2881(08)60129-7
Kolber, Z. Falkowski, P.G. 1993. Use of active fluorescence to estimate phytoplankton photosynthesis in situ. Limnol. Oceanogr. 38(8): 1646-1665. http://dx.doi.org/10.4319/lo.1993.38.8.1646
Kolmogorov A.N. 1941. The local structure of turbulence in incompressible viscous fluid for very large Reynolds numbers. Proc. USSR Acad. Sci. 30: 299-303.
Lande R., Wood M. 1987. Suspension times of particles in the upper ocean. Deep-Sea Res. 34: 61-72. http://dx.doi.org/10.1016/0198-0149(87)90122-1
Lazier J.R.N., Mann K.H. 1989. Turbulence and the diffusive layers around small organisms. Deep-Sea Res. 36: 1721-1733. http://dx.doi.org/10.1016/0198-0149(89)90068-X
Lewis M.R., Horne E.P.W., Cullen J.J., Oakey N.S., Platt T. 1984. Turbulent motions may control phytoplankton photosynthesis in the upper ocean. Nature 311: 49-50. http://dx.doi.org/10.1038/311049a0
Longhurst A., Harrison A. 1989. The biological pump: profiles of plankton production and consumption in the upper ocean. Prog. Oceanogr. 22: 47-123. http://dx.doi.org/10.1016/0079-6611(89)90010-4
Macías D., Lubian L.M., Echevarría F., Huertas E., García C.M. 2008. Chlorophyll maxima and water mass interfaces: tidally induced dynamics in the Strait of Gibraltar. Deep-Sea Res. I 55: 832-846. http://dx.doi.org/10.1016/j.dsr.2008.03.008
Mann K.H., Lazier J.R.N. 1991. Dynamics of marine ecosystems. Biological-Physical Interactions in the Oceans. Blackwell Scientific Publications.
Margalef R. 1978. Life-forms of phytoplankton as survival alternatives in an unstable environment. Oceanol. Acta 1: 493-509.
Mitchell J.G., Yamazaki H., Seuront L., Wolk F., Li H. 2008. Phytoplankton patch patterns: Seascape anatomy in a turbulent ocean. J. Mar. Sys. 69: 247-253. http://dx.doi.org/10.1016/j.jmarsys.2006.01.019
Osborn T.R. 1980. Estimates of the local rate of vertical diffusion from dissipation measurements. J. Phys. Oceanogr. 10: 83-89. http://dx.doi.org/10.1175/1520-0485(1980)010<0083:EOTLRO>2.0.CO;2
Pérez V., Fernández E., Mara-ón E., Morán X.A.G., Zubkov M.V. 2006. Vertical distribution of phytoplankton biomass, production and growth in the Atlantic subtropical gyres. Deep-Sea Res. I 53: 1616-1634. http://dx.doi.org/10.1016/j.dsr.2006.07.008
Pelegrí J.L., Sangrà P. 1998. A mechanism for layer formation in stratified geophysical flows. J. Geophys. Res. 103: 30679-30693. http://dx.doi.org/10.1029/98JC01627
Platt T. 1972. Local phytoplankton abundance and turbulence. J. Plank. Res. 19: 183-187.
Platt T., Sathyendranath S., Caverhill C., Lewis M.R. 1988. Ocean primary production and available light: further algorithms for remote sensing. Deep-Sea Res. 35: 855-879. http://dx.doi.org/10.1016/0198-0149(88)90064-7
Prairie J.C., Franks P.J.S., Jaffe J.S., Doubell M.J., Yamazaki H. 2011. Physical and biological controls of vertical gradients in phytoplankton. Limnol. Oceanogr. Fluids Environ. 1: 75-90. http://dx.doi.org/10.1215/21573698-1267403
Ramirez-Romero, E., Macias, D., Bruno, M., Reyes, E., Navarro, G., Garcia, C.M. 2012. Submesoscale, tidally-induced biogeochemical patterns in the Strait of Gibraltar. Estuar. Coast. Shelf Sci. 101: 24-32. http://dx.doi.org/10.1016/j.ecss.2012.02.010
Riley G.A., Stomnel H., Bumpus D.F. 1949. Quantitative ecology of the plankton of the Western North Atlantic. Bull. Bingharn Oceanogr. Coll. 12: 1-169.
Ross O.N. 2006. Particles in motion: How turbulence affects plankton sedimentation from an oceanic mixed layer. Geophys. Res. Let. 33: L10609.
Rothschild B.J., Osborn T.R. 1988. Small-scale turbulence and plankton contact rates. J. Plank. Res. 10: 465-474. http://dx.doi.org/10.1093/plankt/10.3.465
Ruiz J., García C.M., Rodríguez J. 1996. Sedimentation loss of phytoplankton cells from the mixed layer: effects of turbulence levels. J. Plank. Res. 18: 1727-1734. http://dx.doi.org/10.1093/plankt/18.9.1727
Ruiz J. 1996. The role of turbulence in the sedimentation loss of pelagic aggregates from the mixed layer. J. Mar. Res. 54: 385-406. http://dx.doi.org/10.1357/0022240963213367
Ruiz J., Macías D., Peters F. 2004. Turbulence increases the average settling velocity of phytoplankton cells. Proc. Natl. Acad. Sci. USA 101: 17720-17724. http://dx.doi.org/10.1073/pnas.0401539101
Sangrà, P., Garcia-Muñoz, C., Garcia, C.M., Marrero-Diaz, A., Sobrino, C., Mouriño-Carballido, B., Aguiar-Gonzalez, B., Henriquez-Pastene, C., Rodriguez-Santana, A., Lubian, L.M., Hernandez-Arencibia, M., Hernandez-Leon, S., Vazquez, E., Estrada-Allis, S.N. Coupling between the upper ocean layer variability and size-fractioned phytoplankton in a no nutrient limited environment. Mar. Ecol. Prog. Ser. Submitted
Seuront L., Lagadeuc Y. 1997. Characterization of space-time variability in stratified and mixed coastal waters (Baie des Chaleurs, Quebec, Canada): application of fractal theory. Mar. Ecol. Prog. Ser. 159: 81-95. http://dx.doi.org/10.3354/meps159081
Steele J.H., Yentsch C.S. 1960. The vertical distribution of chlorophyll. J. Exp. Mar. Biol. Ecol. 39: 217-226.
Tennekes H., Lumley J.L. 1972. A First Course in Turbulence. Massachussetts Institute of Technology.
Visser A.W. 1997. Using random walk models to simulate the vertical distribution of particles in a turbulent water column. Mar. Ecol. Progr. Ser. 158: 275-281. http://dx.doi.org/10.3354/meps158275
Wang L.P., Maxey M.R. 1993. Settling velocity and concentration distribution of heavy particles in homogeneous isotropic turbulence.J. Fluid. Mech. 256: 27-68. http://dx.doi.org/10.1017/S0022112093002708
Wolk F., Yamazaki H., Seuront L., Lueck R.G. 2002. A New Free-Fall Profiler for Measuring Biophysical Microstructure. J. Atmos. Ocean Tech. 19: 780-793. http://dx.doi.org/10.1175/1520-0426(2002)019<0780:ANFFPF>2.0.CO;2
Yamazaki H., Mitchell J.G., Seuront L., Wolk F., Li H. 2006. Phytoplankton microstructure in fully developed oceanic turbulence. Geophys. Res. Lett. 33: L01603. http://dx.doi.org/10.1029/2005GL024103
Yamazaki H., Honma H., Nagai T., Doubell M.J., Amakasu K., Kumagai M. 2010. Multilayer biological structure and mixing in the upper water column of Lake Biwa during summer 2008. Limnology 11: 63-70. http://dx.doi.org/10.1007/s10201-009-0288-2
Yang C.Y., Lei U. 1998. The role of the turbulent scales in the settling velocity of heavy particles in homogeneous isotropic turbulence. J. Fluid Mech. 371: 179-205. http://dx.doi.org/10.1017/S0022112098002328
Yang T.S., Shy S.S. 2003. The settling velocity of heavy particles in an aqueous near-isotropic turbulence. Phys. Fluids 15: 868-880. http://dx.doi.org/10.1063/1.1557526
Zhou Q., Cheng N.S. 2009. Experimental investigation of single particle settling in turbulence generated by oscillating grid. Chem. Eng. J. 149: 289-300. http://dx.doi.org/10.1016/j.cej.2008.11.004
Zubkov M.V., Sleigh M.A., Burkill P., Leakey R.J.G. 2000. Picoplankton community structure on the Atlantic Meridional Transect: a comparison between seasons. Prog. Oceanogr. 45: 369-386. http://dx.doi.org/10.1016/S0079-6611(00)00008-2
Descargas
Publicado
Cómo citar
Número
Sección
Licencia
Derechos de autor 2013 Consejo Superior de Investigaciones Científicas (CSIC)
Esta obra está bajo una licencia internacional Creative Commons Atribución 4.0.
© CSIC. Los originales publicados en las ediciones impresa y electrónica de esta Revista son propiedad del Consejo Superior de Investigaciones Científicas, siendo necesario citar la procedencia en cualquier reproducción parcial o total.
Salvo indicación contraria, todos los contenidos de la edición electrónica se distribuyen bajo una licencia de uso y distribución “Creative Commons Reconocimiento 4.0 Internacional ” (CC BY 4.0). Consulte la versión informativa y el texto legal de la licencia. Esta circunstancia ha de hacerse constar expresamente de esta forma cuando sea necesario.
No se autoriza el depósito en repositorios, páginas web personales o similares de cualquier otra versión distinta a la publicada por el editor.
