Physiological acclimation by phytoplankton explains observed changes in Si and N uptake rates during the SERIES iron-enrichment experiment

During the SERIES iron-enrichment experiment in the eastern subarctic Pacific, after addition of iron and its subsequent depletion, the Si:N drawdown ratio increased at approximately the time that diatoms became iron limited. Laboratory studies have reported that this results from a decrease in the rate of N uptake together with a more moderate decrease in the rate of Si uptake for iron-limited cultures compared to iron-replete cultures. However, for SERIES Boyd et al. (Limnol. Oceanogr. 50 (2005)) reported an unexplained increase in the rate of Si uptake at the onset of iron limitation and suggested that studies of nutrient uptake kinetics should be undertaken in search of an explanation. We compare the classic Michealis–Menten (MM) kinetics to the recently developed optimal uptake (OU) kinetics (the SPONGE: Smith and Yamanaka. Limnol. Oceanogr. 52 (2007)) within a variable-composition model, which employs cell quotas for each relevant nutrient, applied to the multi-element (C, N, Si, Fe) dynamics during SERIES. Using the Monte Carlo Markov Chain, we fit two versions of the model (differing only in the equations for nutrient uptake) to the available data for nutrient concentrations, chlorophyll, biogenic silica and particulate organic carbon and specific growth rates.With either uptake kinetics, the model reproduces observed concentrations well for nutrients and somewhat less well for chlorophyll. The different uptake kinetics yield greater differences in modeled elemental composition of phytoplankton and biomass of phytoplankton and zooplankton, which are not directly constrained by data. MM kinetics cannot reproduce the observed increase in Si uptake rate as a function of the decreasing trend in concentration of silicic acid, and it predicts Si limitation throughout nearly all of the experiment after iron-fertilization. In contrast, OU kinetics reproduces the increase in Si uptake rate and matches the observation-based estimate for the timing of the return to iron limitation. The key assumption of the SPONGE, that uptake rates of all nutrients depend on physiological acclimation by phytoplankton as a function of the ambient concentration of the growth-limiting nutrient, was originally formulated for modeling chemostat experiments. We show that it also agrees with the observations from this field experiment and provides an explanation for the increases in Si uptake rate and Si:N drawdown ratio.     

One Dimensional Ecosystem Model Simulation of the Effects of Vertical Dilution by the Winter Mixing on the Spring Diatom Bloom

A one-dimensional ecosystem model has been used to investigate the processes relevant to the spring diatom bloom which play important roles in the biogeochemical cycle in the western subarctic Pacific. The model represents the plankton dynamics and the nutrient cycles in the spring diatom bloom; its results show the importance of dilution by deep mixing in winter. It is supposed that the vertically integrated biomass of phytoplankton decreases in the winter due to the decrease of photosynthesis, because the deep mixing transports phytoplankton to a layer with a low light level. However, the observed integrated diatom biomass increases as the mixed layer deepens. This is because the decrease of concentration due to dilution by mixing causes the diatom grazed pressure to be less significant than diatom photosynthesis. In other words, the effect of dilution on the grazed rate is more significant than the effect on the photosynthesis rate because the grazed rate depends on the concentrations of both diatom and grazer, whereas the photosynthesis rate depends only diatom concentration. The average specific diatom grazed rate, defined as grazed rate divided by diatom biomass, decreases by 35% associated with the deepening, while the average specific photosynthesis rate of diatom decreases by 11%. As a result, the average specific net diatom growth rate during the deep mixing is about 70% of its maximum during the spring diatom bloom. The deep mixing significantly affects the amplitude of the spring diatom bloom not only by the supply of nutrients but also by the dilution which drastically decreases the grazed pressure. This revised version was published online in July 2006 with corrections to the Cover Date.