Oceanography The Official Magazine of
The Oceanography Society
Volume 21 Issue 04

View Issue TOC
Volume 21, No. 4
Pages 46 - 59

New Insights into the Controls and Mechanisms of Plankton Productivity in Coastal Upwelling Waters of the Northern California Current System

Raphael M. Kudela Neil S. Banas John A. BarthElizabeth R. Frame David A. Jay John L. Largier Evelyn J. LessardTawnya D. PetersonAndrea J. Vander Woude
Article Abstract

During the lifetime of the National Science Foundation’s Coastal Ocean Processes program, four experiments were conducted on the US West Coast in the northern California Current System. Although each project had a unique scientific focus, all four addressed the mechanisms causing eastern boundary current systems in general, and the California Current System in particular, to be biologically rich, from phytoplankton to apex predators. Taken together, findings from these projects provide new insights into the canonical view that upwelling systems are simple wind-driven “conveyor belts,” bringing cold, nutrient-rich waters to the well-lit surface ocean where biological organisms flourish. We highlight new insights and advances gained from these programs, including recognition that (a) elements other than nitrogen, particularly iron, may limit the base of the food chain, and (b) the source of these nutrients is not solely a result of wind-driven Ekman transport. The importance of retentive features has clearly emerged, whether these are associated with topography, bathymetry, or more transient features such as river plumes. These new insights into the drivers and fate of this high biological productivity should greatly improve current and future generations of ecosystem models and provide a better understanding of the unique physical-biological coupling that makes the California Current System so rich.


Kudela, R.M., N.S. Banas, J.A. Barth, E.R. Frame, D.A. Jay, J.L. Largier, E.J. Lessard, T.D. Peterson, and A.J. Vander Woude. 2008. New insights into the controls and mechanisms of plankton productivity in coastal upwelling waters of the northern California Current System. Oceanography 21(4):46–59, https://doi.org/10.5670/oceanog.2008.04.


Editor’s Note: Oceanography does not usually permit citation of articles that are in review; however, because of the rapidly advancing nature of this issue’s topics, we are making an exception. Updates on the status of manuscripts cited as in review here will be posted on the CoOP Web site (http://www.skio.usg.edu/coop).

Banas, N.S., P. MacCready, and B.H. Hickey. 2008. The Columbia River plume as cross-shelf exporter and along-coast barrier. Continental Shelf Research doi:10.1016/j.csr.2008.03.011.

Banas, N.S., E.J. Lessard, R.M. Kudela, P. MacCready, T.D. Peterson, B.M. Hickey, and E. Frame. In review. Planktonic growth and grazing in the Columbia River plume region: A biophysical model study.

Barber, R.T., and R.L. Smith. 1981. Coastal upwelling ecosystems. Pp. 31–68 in Analysis of Marine Ecosystems. A.R. Longhurst, ed., Academic Press, New York.

Barth, J.A., and P.A. Wheeler. 2005. Introduction to special section: Coastal advances in shelf transport. Journal of Geophysical Research 110(C10S01): doi:10.1029/2005JC003124.

Barth, J.A., S.D. Pierce, and R.M. Castelao. 2005. Time-dependent, wind-driven flow over a shallow mid-shelf submarine bank. Journal of Geophysical Research 110(C10S05): doi:10.1029/2004JC002761.

Beardsley, R., and S. Lentz. 1987. The coastal ocean dynamics experiment collection: An introduction. Journal of Geophysical Research 92:1,455–1,464.

Botsford, L.W., C.A. Lawrence, E.P. Dever, A. Hastings, and J. Largier. 2003. Wind strength and biological productivity in upwelling systems: An idealized study. Fisheries Oceanography 12:245–259.

Botsford, L.W., C.A. Lawrence, E.P. Dever, A. Hastings, and J.L. Largier. 2006. Effects of variable upwelling on biological productivity on continental shelves in coastal upwelling systems. Deep-Sea Research Part II 53:3,116–3,140.

Brink, K., and T. Cowles. 1991. The coastal transition zone program. Journal of Geophysical Research 96:14,637–14,647.

Bruland, K., E. Rue, and G. Smith. 2001. Iron and macronutrients in California coastal upwelling regimes: Implications for diatom blooms. Limnology and Oceanography 46:1,661–1,674.

Bruland, K.W., M.C. Lohan, A.M. Aguilar-Islas, G.J. Smith, B. Sohst, and A. Baptista. 2008. Factors influencing the chemistry of the near-field Columbia River Plume: Nitrate, silicic acid, dissolved Fe and dissolved Mn. Journal of Geophysical Research doi:10.1029/2007JC004702

Brzezinski, M.A., D.R. Phillips, F.P. Chavez, G.E. Friederich, and R.C. Dugdale. 1997. Silica production in the Monterey, California, upwelling system. Limnology and Oceanography 42:1,694–1,705.

Buck, K.N., M.C. Lohan, C.J.M. Berger, and K.W. Bruland. 2007. Dissolved iron speciation in two distinct river plumes and an estuary: Implications for riverine iron supply. Limnology and Oceanography 52:843–855.

Calbet, A., and M.R. Landry. 2004. Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems. Limnology and Oceanography 49:51–57.

Chase, Z., B. Hales, T. Cowles, R. Schwartz, and A. van Geen. 2005. Distribution and variability of iron input to Oregon coastal waters during the upwelling season. Journal of Geophysical Research 110(C10S12): doi:10.1029/2004JC002590.

Chase, Z., P. Strutton, and B. Hales. 2007. Iron links river runoff and shelf width to phytoplankton biomass along the US west coast. Geophysical Research Letters 34(L04607): doi:10.1029/2006GL028069.

Croll, D.A., B. Marinovic, S. Benson, F.P. Chavez, N. Black, R. Ternullo, and B.R. Tershy. 2005. From wind to whales: Trophic links in a coastal upwelling system. Marine Ecology Progress Series 289:117–130.

Dever, E.P., C.E. Dorman, and J.L. Largier. 2006. Surface boundary-layer variability off Northern California, USA, during upwelling. Deep-Sea Research Part II 53:2,887–2,905.

Dorman, J., S. Bollens, and A. Slaughter. 2005. Population biology of euphausiids off northern California and effects of short time-scale wind events on Euphausia pacifica. Marine Ecology Progress Series 288:183–198.

Dugdale, R., and F. Wilkerson. 1989. New production in the upwelling center off Point Conception, California: Temporal and spatial patterns. Deep-Sea Research 36:985–1,007.

Dugdale, R., F. Wilkerson, V. Hogue, and A. Marchi. 2006. Nutrient controls on new production in the Bodega Bay, California, coastal upwelling plume. Deep-Sea Research Part II 53:3,049–3,062.

Eisner, L., and T. Cowles. 2005. Spatial variations in phytoplankton pigment ratios, optical properties, and environmental gradients in Oregon coast surface waters. Journal of Geophysical Research 110(C10S14): doi:10.1029/2004JC002614.

Enriquez, A.G., and C.A. Friehe. 1995. Effects of wind stress and wind stress curl variability on coastal upwelling. Journal of Physical Oceanography 25:1,651–1,671.

FAO. 2002. The state of world fisheries and aquaculture 2002. FAO, Rome, Italy.

Frame, E.R., and E.J. Lessard. In review. Does the Columbia River Plume influence phytoplankton community structure along the Washington and Oregon coasts?

Fung, Y.I., S.K. Meyn, I. Tegen, S.C. Doney, J.G. John, and J.K.B. Bishop. 2000. Iron supply and demand in the upper ocean. Global Biogeochemical Cycles 14:281–295.

Gan, J., and J.S. Allen. 2002. A modeling study of shelf circulation off northern California in the region of the Coastal Ocean Dynamics Experiment: Response to relaxation of upwelling winds. Journal of Geophysical Research 107(C9):3123: doi:10.1029/2000JC000768.

Graham, W.M., and J.L. Largier. 1997. Upwelling shadows as nearshore retention sites: The example of northern Monterey Bay. Continental Shelf Research 17:509–532.

Hales, B., J. Moum, P. Covert, and A. Perlin. 2005a. Irreversible nitrate fluxes due to turbulent mixing in a coastal upwelling system. Journal of Geophysical Research 110(C10511): doi:10.1029/2004JC002685.

Hales, B., T. Takahashi, and L. Bandstra. 2005b. Atmospheric CO2 uptake by a coastal upwelling system. Global Biogeochemical Cycles 19(GB1009): doi:10.1029/2004GB002295.

Hurst, M.P., and K.W. Bruland. 2007. The effect of the San Francisco Bay plume on trace metal and nutrient distributions in the Gulf of the Farallones. Geochimica et Cosmochimica Acta 72:395–411.

Hutchins, D., and K. Bruland. 1998. Iron-limited diatom growth and Si:N uptake ratios in a coastal upwelling regime. Nature 393:561–564.

Hutchins, D., G. DiTullio, Y. Zhang, and K. Bruland. 1998. An iron limitation mosaic in the California upwelling regime. Limnology and Oceanography 43:1,037–1,054.

Huyer, A., J. Fleischbein, J. Keister, P. Kosro, N. Perlin, R. Smith, and P. Wheeler. 2005. Two coastal upwelling domains in the northern California Current system. Journal of Marine Research 63:901–929.

Jay, D.A., J. Pan, P.M. Orton, and A. Horner-Devine. In press. Asymmetry of tidal plume fronts in an eastern boundary current regime. Journal of Marine Systems.

Johnson, K.S., F.P. Chavez, V.A. Elrod, S.E. Fitzwater, J.T. Pennington, K.R. Buck, and P.M. Walz. 2001. The annual cycle of iron and the biological response in central California coastal waters. Geophysical Research Letters 28:1,247–1,250.

Kaplan, D.M., and J.L. Largier. 2006. HF-radar-derived origin and destination of surface waters off Bodega Bay, California. Deep Sea Research Part II 53:2,906–2,930.

King, A.L., and K. Barbeau. 2007. Evidence for phytoplankton iron limitation in the southern California current system. Marine Ecology Progress Series 342:91–104.

Kudela, R.M., W.P. Cochlan, and R.C. Dugdale. 1997. Carbon and nitrogen uptake response to light by phytoplankton during an upwelling event. Journal of Plankton Research 19:609–630.

Kudela, R., N. Garfield, and K. Bruland. 2006. Bio-optical signatures and biogeochemistry from intense upwelling and relaxation in coastal California. Deep-Sea Research Part II 53:2,999–3,022.

Kudela, R.M., and T.D. Peterson. In review. Influence of a buoyant river plume on phytoplankton nutrient dynamics: What controls standing stocks and productivity? 

Lamb, J., and W. Peterson. 2005. Ecological zonation of zooplankton in the COAST study region off central Oregon in June and August 2001 with consideration of retention mechanisms. Journal of Geophysical Research 110(C10S15): doi:10.1029/2004JC002520.

Largier, J. 2004. The importance of retention zones in the dispersal of larvae. American Fisheries Society Symposium 45:105–122.

Largier, J., C. Lawrence, M. Roughan, D. Kaplan, E. Dever, C. Dorman, R. Kudela, S. Bollens, F. Wilkerson, R. Dugdale, and others. 2006. WEST: A northern California study of the role of wind-driven transport in the productivity of coastal plankton communities. Deep-Sea Research Part II 53:2,833–2,849.

Lassiter, A., F. Wilkerson, R. Dugdale, and V. Hogue. 2006. Phytoplankton assemblages in the CoOP-WEST coastal upwelling area. Deep-Sea Research Part II 53:3,063–3,077.

Lessard, E.J., and E.R. Frame. 2008. The influence of the Columbia River Plume on patterns of phytoplankton growth, grazing and chlorophyll on the Washington and Oregon coasts. Paper presented at the 2008 Ocean Sciences Meeting, AGU/ASLO/TOS/ERF, Orlando, FL.

Mackas, D., W. Peterson, M. Ohman, and B. Lavaniegos. 2006. Zooplankton anomalies in the California Current system before and during the warm ocean conditions of 2005. Geophysical Research Letters 33(L22S07): doi:10.1029/2006GL027930.

Martin, J., and S. Fitzwater. 1988. Iron deficiency limits phytoplankton growth in the northeast Pacific subarctic. Nature 331:341–343.

MacCready, P., N.S. Banas, B.M. Hickey, E.P. Dever, and Y. Liu. 2008. A model study of tide- and wind-induced mixing in the Columbia River estuary and plume. Continental Shelf Research: doi:10.1016/j.csr.2008.03.015.

Munchow, A. 2000. Wind stress curl forcing of the coastal ocean near Point Conception, California. Journal of Physical Oceanography 30:1,265–1,280.

Nash, J., and J. Moum. 2005. River plumes as a source of large-amplitude internal waves in the coastal ocean. Nature 437:400–403: doi:10.1038/nature03936.

Nicholson, D., S. Dyhrman, F. Chavez, and A. Paytan. 2006. Alkaline phosphatase activity in the phytoplankton communities of Monterey Bay and San Francisco Bay. Limnology and Oceanography 51:874–883.

Oke, P.R., J.S. Allen, R.N. Miller, and G.D. Egbert. 2002. A modeling study of the three-dimensional continental shelf off Oregon. Part II: Dynamical analysis. Journal of Physical Oceanography 32:1,383–1,403.

Orton, P.M., and D.A. Jay. 2005. Observations at the tidal plume front of a high-volume river outflow. Geophysical Research Letters 32(L11605): doi:10.1029/2005GL022372.

Pan, J., and D.A. Jay. 2008. Dynamical characteristics and horizontal transport of large amplitude internal solitons generated at the Columbia River plume front. Continental Shelf Research 28: doi:10.1016/ j.csr.2008.01.002.

Peterson, J., and W. Peterson. In review. Influence of the Columbia River plume on the cross-shelf transport of zooplankton.

Ruttenberg, K., and S. Dyhrman. 2005. Temporal and spatial variability of dissolved organic and inorganic phosphorous, and metrics of phosphorous bioavailability in an upwelling-dominated coastal system. Journal of Geophysical Research 110(C10S13): doi:10.1029/2004JC002837.

Ryther, J., and W. Dunstan. 1971. Nitrogen, phosphorous, and eutrophication in the coastal marine environment. Science 171:1,008–1,013.

Send, U., R.C. Beardsley, and C.D. Winant. 1987. Relaxation from upwelling in the Coastal Ocean Dynamics Experiment. Journal of Geophysical Research 92:1,683–1,698.

Sherr, E., B. Sherr, and P. Wheeler. 2005. Distribution of coccoid cyanobacteria and small eukaryotic phytoplankton in the upwelling system off the Oregon coast during 2001 and 2002. Deep-Sea Research Part II 52:317–330.

Slaughter, A., S. Bollens, and G. Rollwagen Bollens. 2006. Grazing impact of mesozooplankton in an upwelling region off northern California, 2000–2003. Deep-Sea Research Part II 53:3,099–3,115.

Sydeman, W., R. Bradley, P. Warzybok, C. Abraham, J. Jahncke, K. Hyrenbach, V. Kousky, J. Hipfner, and M. Ohman. 2006. Planktivorous auklet Ptychoramphus aleuticus responses to ocean climate, 2005: Unusual atmospheric blocking? Geophysical Research Letters 33(L22S09): doi:10.1029/2006GL026736.

Vander Woude, A. 2006. Coastal retentive embayments north and south of Point Reyes, California: Existence, time scales and carbon significance. PhD Dissertation, University of California, Santa Cruz.

Vander Woude, A., J. Largier, and R. Kudela. 2006. Nearshore retention of upwelled waters north and south of Point Reyes (northern California)—Patterns of surface temperature and chlorophyll observed in CoOP WEST. Deep-Sea Research Part II 53:2,985–2,998.

Wetz, M., and P. Wheeler. 2003. Production and partitioning of organic matter during simulated phytoplankton blooms. Limnology and Oceanography 48:1,808–1,817.

Ware, D., and R. Thomson. 2005. Bottom-up ecosystem trophic dynamics determine fish production in the Northeast Pacific. Science 308:1,280–1,284.

Weise, M., D. Costa, and R. Kudela. 2006. Movement and diving behavior of male California sea lion (Zalophus californianus) during anomalous oceanographic conditions of 2005 compared to those of 2004. Geophysical Research Letters 33(L22S10): doi:10.1029/2006GL027113.

Wilkerson, F., A. Lassiter, R. Dugdale, A. Marchi, and V. Hogue. 2006. The phytoplankton bloom response to wind events and upwelled nutrients during the CoOP-WEST study. Deep-Sea Research Part II 53:3,023–3,048.

Wing, S.R., J.L. Largier, L.W. Botsford, and J.F. Quinn. 1995a. Settlement and transport of benthic invertebrates in an intermittent upwelling region. Limnology and Oceanography 40:316–329.

Wing, S.R., L.W. Botsford, J.L. Largier, and L.E. Morgan. 1995b. Spatial structure of relaxation events and crab settlement in the northern California upwelling region. Marine Ecology Progress Series 128:199–211.

Wing, S.R., L.W. Botsford, S.V. Ralston, and J.L. Largier. 1998. Meroplanktonic distribution and circulation in a coastal retention zone of the northern California upwelling system. Limnology and Oceanography 43:1,710–1,721.

Zaron, E.D., and D.A. Jay. In review. Mixing in the tidal plume of the Columbia River.