Origin and maintenance of a high nitrate condition in the equatorial Pacific

Abstract The physical and biological causes for the equatorial nutrient anomaly were investigated using an ecosystem model embedded within an ocean general circulation model to determine the nitrate budget for the equatorial Pacific Ocean. In the 140°W region the effects of mixing on nitrate concentration were small compared to the effects of advection: upwelling and zonal transport to the east in the Equatorial Undercurrent were the major processes in the nitrate budget. At 140°W on the equator annual JNO3, the total net physical supply of nitrate to the euphotic layer, was 3.76 mmol m−2 day−1; the vertical integrated (0–120m) new production calculated from the ecosystem model was 3.36 mmol m−2 day−1 or, in carbon units, 22.26 mmol C m−2 day−1. The vertical supply of nitrate (−w∂NO3/∂z) due to the upwelling is controlled by two factors, the vertical velocity and vertical gradient of nitrate concentration. The vertical velocity reaches the maximum during climatological fall, but the vertical gradient of nitrate is weaker in the fall. Therefore, the vertical supply of nitrate is smaller than in spring. To investigate the role of physiological limitation of phytoplankton photosynthesis and specific growth rate on the maintenance of the high nutrient-low chlorophyll (HNLC) condition, a model experiment was performed that included, unchanged from previous model runs, the physical conditions and density-dependent grazing function, but greatly reduced physiological limitations by increasing α (initial slope of P-I curve) and Pmax (maximum specific growth rate) values. When this was done, vertical integrated primary production at 140°W on the equator doubled (from 83 to 166 mmol C m−2 day−1), but the zooplankton grazing on the phytoplankton also doubled (from 75 to 150 mmol C m−2 day−1). Zooplankton biomass doubled, but there was only a slight increase in phytoplankton biomass; no phytoplankton bloom formed in this model experiment. With potential physiological limitations of phytoplankton rates greatly reduced, the characteristic equatorial plume of unused surface layer nitrate still persisted; but the nitrate-rich plume was smaller in horizontal extent and the maximum concentration was reduced by half from observed concentrations. While the reduction in the extent of the nitrate-rich plume indicates that physiological limitation plays a significant role in the maintenance of the nutrient anomaly, its persistence demonstrates that physical processes and grazing also are involved.

[1]  K. Coale,et al.  Iron distributions in the equatorial Pacific: Implications for new production , 1997 .

[2]  T. Platt,et al.  Vertical Nitrate Fluxes in the Oligotrophic Ocean , 1986, Science.

[3]  F. Morel,et al.  The equatorial Pacific Ocean: Grazer-controlled phytoplankton populations in an iron-limited ecosystem1 , 1994 .

[4]  D. F. Winter,et al.  A theoretical study of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern U.S. coast , 1977 .

[5]  R. Feely,et al.  Physical and Biological Controls on Carbon Cycling in the Equatorial Pacific , 1994, Science.

[6]  John H. Martin glacial-interglacial Co2 change : the iron hypothesis , 1990 .

[7]  D. Halpern,et al.  Vertical motion in the upper ocean of the equatorial Eastern Pacific , 1987 .

[8]  Daniel Kamykowski,et al.  Predicting plant nutrient concentrations from temperature and sigma-t in the upper kilometer of the world ocean , 1986 .

[9]  B. Frost The role of grazing in nutrient-rich areas of the open sea , 1991 .

[10]  R. Bidigare,et al.  Phytoplankton photosynthesis parameters along 140°W in the equatorial Pacific , 1995 .

[11]  F. Chavez,et al.  Estimating new production in the equatorial Pacific Ocean at 150°W , 1992 .

[12]  B. Frost,et al.  Grazing control of phytoplankton stock in the open subarctic Pacific Ocean: a model assessing the role of mesozooplankton, particularly the large calanoid copepods Neocalanus spp. , 1987 .

[13]  B. Peterson,et al.  Particulate organic matter flux and planktonic new production in the deep ocean , 1979, Nature.

[14]  Francisco P. Chavez,et al.  Standing stocks of particulate carbon and nitrogen in the equatorial Pacific at 150°W , 1992 .

[15]  James W. Murray,et al.  A U.S. JGOFS process study in the equatorial Pacific (EqPac): Introduction , 1995 .

[16]  R. Barber Introduction to the WEC88 cruise: An investigation into why the equator is not greener , 1992 .

[17]  P. Buat-Ménard The role of air-sea exchange in geochemical cycling , 1986 .

[18]  R. Duce THE IMPACT OF ATMOSPHERIC NITROGEN, PHOSPHORUS, AND IRON SPECIES ON , 1986 .

[19]  W. Richard,et al.  TEMPERATURE AND PHYTOPLANKTON GROWTH IN THE SEA , 1972 .

[20]  S. Philander,et al.  Simulation of El Niño of 1982–1983 , 1985 .

[21]  H. Bryden,et al.  Diagnostic Model of the Three-Dimensional Circulation in the Upper Equatorial Pacific Ocean , 1985 .

[22]  H. Ducklow,et al.  A nitrogen-based model of plankton dynamics in the oceanic mixed layer , 1990 .

[23]  J. Goering,et al.  UPTAKE OF NEW AND REGENERATED FORMS OF NITROGEN IN PRIMARY PRODUCTIVITY1 , 1967 .

[24]  R. Pacanowski,et al.  Parameterization of Vertical Mixing in Numerical Models of Tropical Oceans , 1981 .

[25]  M. Fasham Variations in the seasonal cycle of biological production in subarctic oceans: A model sensitivity analysis , 1995 .

[26]  Richard W. Reynolds,et al.  A Real-Time Global Sea Surface Temperature Analysis , 1988 .

[27]  D. Halpern A pacific equatorial temperature section from 172°E to 110°W during winter and spring 1979 , 1980 .

[28]  F. Chavez,et al.  Growth rates, grazing, sinking, and iron limitation of equatorial Pacific phytoplankton , 1991 .

[29]  S. Levitus Climatological Atlas of the World Ocean , 1982 .

[30]  T. Ku,et al.  228Ra-derived nutrient budgets in the upper equatorial Pacific and the role of 'new' silicate in limiting productivity , 1995 .

[31]  S. Levitus,et al.  Distribution of nitrate, phosphate and silicate in the world oceans , 1993 .

[32]  S. Fitzwater,et al.  The case for iron , 1991 .

[33]  J. Toggweiler,et al.  A seasonal three‐dimensional ecosystem model of nitrogen cycling in the North Atlantic Euphotic Zone , 1993 .

[34]  R. Reynolds,et al.  Historical trends in the surface temperature over the oceans based on the COADS , 1987 .

[35]  H. Jannasch,et al.  Nutrient assimilation, export production and 234Th scavenging in the eastern equatorial Pacific , 1989 .

[36]  Curtiss O. Davis,et al.  Photosynthetic characteristics and estimated growth rates indicate grazing is the proximate control of primary production in the equatorial Pacific , 1992 .

[37]  R. Harris,et al.  Feeding, Growth and Reproduction of the Marine Planktonic Copepod Pseudo-Calanus Elongatus Boeck , 1976, Journal of the Marine Biological Association of the United Kingdom.

[38]  F. Chavez,et al.  An estimate of new production in the equatorial Pacific , 1987 .

[39]  F. Chai,et al.  Primary productivity and its regulation in the equatorial Pacific during and following the 1991–1992 El Niño , 1996 .

[40]  J. Ryther,et al.  Organic chelators: Factors affecting primary production in the cromwell current upwelling☆ , 1969 .

[41]  B. Frost,et al.  Grazing and iron limitation in the control of phytoplankton stock and nutrient concentration: a chemostat analogue of the Pacific equatorial upwelling zone , 1992 .

[42]  J. Nihoul Coupled ocean-atmosphere models , 1985 .

[43]  W. Jenkins,et al.  Nitrate flux into the euphotic zone near Bermuda , 1988, Nature.

[44]  W. G. Harrison,et al.  Primary productivity and size structure of phytoplankton biomass on a transect of the equator at 135°W in the Pacific Ocean , 1990 .

[45]  D. F. Winter,et al.  Sensitivity analysis of a mathematical model of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern U.S. coast , 1979 .

[46]  K. Wyrtki An Estimate of Equatorial Upwelling in the Pacific , 1981 .

[47]  W. Hurlin,et al.  Simulation of the Seasonal Cycle of the Tropical Pacific Ocean , 1987 .

[48]  A. Gargett Physical processes and the maintenance of nutrient‐rich euphotic zones , 1991 .

[49]  A. J. Watson,et al.  Testing the iron hypothesis in ecosystems of the equatorial Pacific Ocean , 1994, Nature.

[50]  J. McCarthy,et al.  New production along 140°W in the equatorial Pacific during and following the 1992 El Niño event , 1996 .

[51]  C. Davis,et al.  Primary production estimates from recordings of solar-stimulated fluorescence in the equatorial Pacific at 150°W , 1992 .

[52]  J. Steele,et al.  The role of predation in plankton models , 1992 .

[53]  M. Cox A primitive equation, 3-dimensional model of the ocean , 1984 .

[54]  James J. McCarthy,et al.  HALF‐SATURATION CONSTANTS FOR UPTAKE OF NITRATE AND AMMONIUM BY MARINE PHYTOPLANKTON1 , 1969 .

[55]  P. Liss,et al.  The Role of Episodic Atmospheric Nutrient Inputs in the Chemical and Biological Dynamics of Oceanic Ecosystems , 1991 .

[56]  R. Duce,et al.  Atmospheric transport of iron and its deposition in the ocean , 1991 .

[57]  F. Chavez,et al.  Biological Consequences of El Ni�o , 1983, Science.

[58]  David M. Karl,et al.  VERTEX: carbon cycling in the northeast Pacific , 1987 .

[59]  T. Packard,et al.  Productivity in upwelling areas deduced from hydrographic and chemical fields1 , 1986 .

[60]  Michael R. Landry,et al.  Microzooplankton grazing in the central equatorial Pacific during February and August, 1992 , 1995 .

[61]  H. Sverdrup,et al.  On Conditions for the Vernal Blooming of Phytoplankton , 1953 .

[62]  John J. Walsh,et al.  Herbivory as a factor in patterns of nutrient utilization in the sea1 , 1976 .

[63]  Francisco P. Chavez,et al.  Regulation of primary productivity rate in the equatorial Pacific , 1991 .