Importance of microzooplankton for sustaining high mesozooplankton biomass during post-bloom period in the Oyashio region of the western subarctic Pacific

We investigated the plankton community structure and biomass during the post-bloom season in the Oyashio region of the western subarctic Pacific, including pico-, nano-, microplankton and mesozooplankton. We found that the nitrate, phosphate and silicic acid concentrations remained high at >4.2 μM, >0.77 μM and >7.1 μM, respectively, in the euphotic layer at almost all sampling stations, but that the chlorophyll a concentrations were low (<3 µg Chl. a l−1). These findings indicate high nutrient and low chlorophyll (HNLC)-like conditions. In the phytoplankton community, pennate diatoms, the larger subpopulation of pico-sized eukaryotic phytoplankton, and nano-flagellates substantially contributed to the low biomass of the chain-forming centric diatoms that mainly comprised the spring phytoplankton bloom. The microzooplankton biomass was 2.7–4.4 fold greater than the phytoplankton biomass in the surface layer. Naked ciliates substantially contributed to the microzooplankton community (40 %–87 %). The naked ciliate growth rates during our in situ bottle incubation experiments were significantly greater than the maximum growth rates as calculated from cell volume and water temperature. The mesozooplankton biomass was mainly composed of krill and copepods and was 5.9–9.3 fold higher than the microzooplankton biomass. This inverted biomass pyramid with relatively low microzooplankton and high mesozooplankton biomass may be explained by the high production and growth rates of the microzooplankton. The ratio of phytoplankton growth (µ, d−1) to grazing mortality (m, d−1) by microzooplankton were relatively low at 0.26–0.44 m/µ in our dilution experiments. These low values indicate that microzooplankton grazing does not regulate phytoplankton growth and suggests that microzooplankton feed on an alternative nutritional source, such as heterotrophic prey items, or mixotrophy to fulfill their growth needs. Additional research is needed during the post-bloom period to further evaluate the mechanisms that sustain microzooplankton dominance and production in the Oyashio region under the HNLC-like conditions, especially for naked ciliates.