An ecosystem modeling study of spatio-temporal variations of phytoplankton distribution in the Okhotsk Sea

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Abstract

A three-dimensional ecosystem–physical-coupled model is applied to the Okhotsk Sea. Factors that determine the spatial distribution of phytoplankton in the Okhotsk Sea in autumn and spring are analyzed, and the effects of sea ice on the spring bloom are also discussed. One of the most important factors determining the spatial distribution of phytoplankton in autumn is regional variation in mixed layer depth. The model can explain the spatio-temporal variation of chlorophyll-a concentration in the Okhotsk Sea during the spring blooms in 1997 and 2001. The start of the spring bloom in the Okhotsk Sea depends on the light environment. By controlling the light intensity in sea surface water, sea ice controls the timing of the spring bloom.

Introduction

The Okhotsk Sea is one of the most biologically productive regions in the world, and it supports high fisheries production (Fig. 1). It has been reported that a major fraction of the phytoplankton in this sea are diatoms (e.g., Hanzawa et al., 1981). Previous observations revealed maximum diatom cell numbers in spring and minima in autumn (Ohwada, 1957; Hanzawa et al., 1981). Saitoh et al. (1996) showed that the spring bloom was formed between April and May based on the monthly coastal zone color scanner (CZCS)-chlorophyll (Chl) imagery from 1978 until 1986. Shiomoto et al. (1998) suggested that the Chl-a concentration is lower (0–3 mg/m3) in autumn than during the spring bloom (2–10 mg/m3; Nishihama et al., 1989), and that grazing by macrozooplankton (mainly Copepods) may contribute to variations in Chl-a concentration.

The Okhotsk Sea is well known as one of the southern most seasonal sea ice zones in the Northern Hemisphere. In winter, sea ice formation begins around Shantarsky Bay at the end of November, and sea ice extension reaches its maximum in late February or March. Most of the sea ice disappears by May. Sea ice in the Okhotsk Sea is generally advected southward by the prevailing northerly or northwesterly winds. In the southwestern part of the sea, thick “first-year” ice is primarily advected by the East Sakhalin Current (ESC). Some of the ice is advected toward the offshore warm region and melted, making the surface layer water fresher. Some of this water is then frozen again by cooling. This process leads to formation of “new ice” at ice edges (Ohsima et al., 2001). Therefore, the seasonal change of sea ice volume depends on climatic conditions, and has large interannual variations. The sea ice has been considered to play an important role in the high production at the ice edge in the Okhotsk Sea. A large number of studies paid attention to the spring bloom after the sea ice melting (e.g., at the Bering Sea; Niebauer and Alexnder, 1985). However, there are a few studies (e.g., Matsumoto et al., 2004) that discuss the effect of sea ice on the spring bloom in the Okhotsk Sea.

Recently, numerical models have been applied to examine what controls phytoplankton blooms in the Northern Pacific. Kawamiya et al., 1995, Kawamiya et al., 1997 developed a vertical one-dimensional ecological–physical-coupled model and explained the role of zooplankton grazing pressure in the HNLC at OS Papa. Since then, KKYS (the ecological part of their model) has been applied to several different oceanic regimes, the Bermuda Atlantic Time-Series (BATS) site and Japanese coastal regions. The series of studies showed that with changes in only a few parameters, the model could reproduce nitrate and Chl concentrations in those regimes. KKYS is a simple ecological model, composed of six compartments. Based on KKYS, Kishi et al. (2001) developed an eight-compartment ecological model by dividing phytoplankton and zooplankton each into two size categories. Moreover, building upon their model, North Pacific Ecosystem Model Understanding Regional Oceanography (NEMURO) was developed by the PICES CCCC-MODEL TASK TEAM (Eslinger et al., 2000) during a workshop held in January 2000, in NEMURO, Hokkaido, Japan. NEMURO was developed to simulate the lower trophic ecosystem of the Northern Pacific Ocean. Fujii et al. (2002) showed that NEMURO can reproduce the transition of the seasonal dominant species at station KNOT (44°N, 155°E) in their one-dimensional model.

A few models have been made and applied to algal bloom under sea ice. Fennel et al. (2003) made an ecosystem model on seasonal ice zone of the Southern Ocean. In their model, the effects of the flux of light under sea ice, which is used for photosynthesis, and momentum through sea ice, which is used as physical forcing as a boundary condition, are included. But they do not take into consideration the time-dependent change of the thickness of sea ice, nor do they focus on the effects of sea ice on photosynthesis of planktons.

The purposes of this paper are to examine the factors that control a phytoplankton growth in autumn and spring, and to discuss the effect of sea ice on the spring bloom in the Okhotsk Sea with the aid of the ecosystem model. In 2.1 Modeling the autumn bloom, 3.1 Autumn bloom, we present the application of NEMURO coupled with a three-dimensional physical model to the Okhotsk Sea in autumn. We focused on the factors that determine the spatial distribution of phytoplankton in the Okhotsk Sea in autumn. In 2.2 Modeling the spring bloom, 3.2 Spring bloom, we present a modified model including sea ice (including its temporal changes) and discuss the effects of sea ice on the spring bloom, paying attention to the factors that determine the spatial distribution of phytoplankton in the Okhotsk Sea in spring.

Section snippets

Ecosystem–physical-coupled model

A sigma-coordinate modular ocean model (MOM) ver. 1 modified by Harvard University (Harvard Ocean Prediction System, Lozano et al., 1994; Robinson, 1996) is used for the physical model. Pacanowski and Philander's (1981) parameterization is adopted as vertical diffusivity and viscosity. The model domain extends from 42°N to 64°N and from 138°E to 164°E. The horizontal grid scale is 1/4°. The Soya straits are closed, and grids shallower than 50 m depth are masked. There are 15 vertical levels. The

Autumn bloom

The significance of the ESC, which has been presented by schematic figures (e.g., Watanabe, 1963), is not clear in the initial current field (Fig. 3(A)). Also, after 30 days calculation, the ESC of the Okhotsk Sea is not reproduced (Fig. 3(B)). The calculated spatial variations of Chl-a concentration (PS+PL) in the surface layer averaged for 30 days are shown in Fig. 6. Relatively high Chl-a concentration is found in the western Okhotsk Sea. The observed surface Chl-a concentration is 0.72–2.78

Autumn bloom

Observed surface Chl-a concentrations are high in the western part of the Okhotsk Sea in 1996, October–November (Fig. 6). Also, surface Chl-a concentration obtained from monthly mean CZCS-Chl images in October and November (Saitoh et al., 1996) shows that Chl-a concentration is high to the east of Sakhalin Island and low in the Kuril Basin. The simulated Chl-a concentration pattern is similar to the observations (Fig. 6) and satellite imagery (Saitoh et al., 1996). That is, sea surface Chl-a

Conclusion

The factors that determine the spatial distribution of phytoplankton in autumn and spring, and the effects of sea ice on the spring bloom in the Okhotsk Sea were examined using a three-dimensional ecosystem–physical-coupled model.

In autumn, a very shallow mixed layer developed to the east of Sakhalin Island, because the salinity of the surface water was reduced by the discharge from the Amur River. This caused high phytoplankton concentrations there. On the other hand, in the Central Okhotsk

Acknowledgements

We would like to thank Dr. Oshima of Hokkaido University for his fruitful discussions. We wish to express our thanks to Dr. S. Lan Smith of Frontier Research Center for Global Change for his valuable comments and his help in proof reading the manuscript. We thank Dr. Tateyama of Hokkaido University for providing the thickness of sea ice data set. We also thank Prof. Saitoh and Miss Matsumoto of Hokkaido University for providing the OCTS-Chl images. We also appreciate the valuable comments of

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