The significance of phytoplankton photo-adaptation and benthic-pelagic coupling to primary production in the South China Sea: Observations and numerical investigations

The primary production in the South China Sea (SCS) has been assessed by a coupled physical-biogeochemical model with a simple NPZD ecosystem [Liu et al., 2002. Monsoon-forced chlorophyll distribution and primary production in the SCS: observations and a numerical study. Deep-Sea Research I 49(8), 1387-1412]. In recent years there have been an increasing number of observations in the SCS that may be used to check the validity of the previous approach. The coupled model of the SCS mentioned above employs a photo-adaptation scheme for the phytoplankton growth and uses the simplest bottom boundary condition of an inert benthic layer. These adopted schemes are checked against observations at the South-East Asian Time-series Study (SEATS) Station in the northern SCS and in the Gulf of Thailand. Numerical experiments with or without photo-adaptation or active benthic processes are carried out in this study. Additional experiments are performed with different parameters used for these processes. The observations at the SEATS Station provide direct evidence for the variable chlorophyll-to-nitrogen ratio in phytoplankton as required by photo-adaptation. It is concluded that a photo-adaptation scheme is critical to the phytoplankton growth, especially for the development of the subsurface chlorophyll maximum (SCM). Without photo-adaptation, the average value of the vertically integrated primary production (IPP) over the whole SCS domain would be 35% lower. It is noted that, the modeled SCM occurs at depths shallower than observations due to physical as well as biological processes employed by the model. Increasing the upper limit of the chlorophyll-to-nitrogen ratio, as suggested by observations, enhances chlorophyll level in the lower part of the euphotic zone and raises primary productivity in areas with rich nutrient supply. The observed values of the IPP in the Gulf of Thailand clearly demonstrate the importance of the benthic-pelagic coupling to the nutrient cycle. Without benthic nutrient regeneration the model grossly underestimates primary production due to failure to build up the nutrient reserve in the Gulf. On the other hand, a fully regenerated flux of particulate organic nitrogen at the sea floor without denitrification produces too strong a primary production. The improved model uses a higher upper limit for the chlorophyll-to-phytoplankton ratio of 3.5 g Chl/mol N and adopts benthic processes of a coupled nitrification-denitrification scheme with denitrification consuming 14% of the detritus flux at the bottom. The model predicts a mean annual IPP value of 406 mg C m−2 d−1 for the SCS, which is 44% higher than that predicted by the original model. The increase can be broken down to 39% attributed to the benthic nutrient regeneration and 5% to the enhanced photo-adaptation. The average IPP is 390 mg C m−2 d−1 for the basin region (>200 m) and 429 mg C m−2 d−1 for the shelf region (<200 m), both of which compare favorably with observed mean values. The model also predicts a mean nitrogen removal flux of 0.16 mmol N m−2 d−1 during denitrification for the shelf region.