The PICES CCCC (North Pacific Marine Science Organization, ClimateChange and Carrying Capacity program) MODEL Task Team achieved a consensus on the structure of a prototype lower trophic level ecosystem model for the North Pacific Ocean, and named it the North Pacific Ecosystem Model for Understanding Regional Oceanography, "NEMURO". Through an extensive dialog between modelers, plankton biologists and oceanographers, an extensive review was conducted to define NEMURO's process equations and their parameter values for distinct geographic regions. We present in this paper the formulation, structure and governing equations of NEMURO as well as examples to illustrate its behavior. NEMURO has eleven state variables: nitrate, ammonium, small and large phytoplankton biomass, small, large and predatory zooplankton biomass, particulate and dissolved organic nitrogen, particulate silica, and silicic acid concentration. Several applications reported in this issue of Ecological Modelling have successfully used NEMURO, and an extension that includes fish as an additional state variable. Applications include studies of the biogeochemistry of the North Pacific, and variations of its ecosystem's lower trophic levels and two target fish species at regional and basin-scale levels, and on time scales from seasonal to interdecadal.5

Plankton populations in Prince William Sound, Alaska, exhibited pronounced seasonal, annual and longer‐period variability in composition and standing stock in response to physically influenced differences in nutrient availability, and possibly currents that modify local biomass by exchanges with water from the bordering Gulf of Alaska. During springs in which early, strong physical stratification developed, intense, short‐lived phytoplankton blooms occurred. These blooms had relatively short residence times in the water column. In contrast, during springs in which slower, weaker stratification developed, phytoplankton blooms were prolonged and took longer to peak. These slower blooms prolonged the period of phytoplankton production, prolonged interaction with the springtime grazing community and led to the incorporation of more organic matter into pelagic food webs. A coupled biological‐physical simulation of plankton production was used to examine the implications of seasonally varying air and mixed‐layer temperatures, surface winds and incident light on the timing, duration, annual production and standing stock of plankton. Our modelling results reproduced the observed characteristics of the springtime production cycle, and the magnitude of zooplankton stocks for the period 1992–97 but not for 1981–91. These results suggest that for most of the 1990s, bottom‐up influences on nutrient supplies controlled levels of primary consumers, whereas for the 11 years before that, other unknown factors dominated this process. We present the results of a comprehensive, multiyear study of relationships between plankton and physical limitations, and a retrospective analysis of earlier conditions to explore the possible causes for these differences.

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Five years of field, laboratory, and numerical modelling studies demonstrated ecosystem‐level mechanisms influencing the mortality of juvenile pink salmon and Pacific herring. Both species are prey for other fishes, seabirds, and marine mammals in Prince William Sound. We identified critical time‐space linkages between the juvenile stages of pink salmon and herring rearing in shallow‐water nursery areas and seasonally varying ocean state, the availability of appropriate zooplankton forage, and the kinds and numbers of predators. These relationships defined unique habitat dependencies for juveniles whose survivals were strongly linked to growth rates, energy reserves, and seasonal trophic sheltering from predators. We found that juvenile herring were subject to substantial starvation losses during a winter period of plankton diminishment, and that predation on juvenile pink salmon was closely linked to the availability of alternative prey for fish and bird predators. Our collaborative study further revealed that juvenile pink salmon and age‐0 herring exploit very different portions of the annual production cycle. Juvenile pink salmon targeted the cool‐water, early spring plankton bloom dominated by diatoms and large calanoid copepods, whereas young‐of‐the‐year juvenile herring were dependent on warmer conditions occurring later in the postbloom summer and fall when zooplankton was composed of smaller calanoids and a diversity of other taxa. The synopsis of our studies presented in this volume speaks to contemporary issues facing investigators of fish ecosystems, including juvenile fishes, and offers new insight into problems of bottom‐up and top‐down control. In aggregate, our results point to the importance of seeking mechanistic rather than correlative understandings of complex natural systems.

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